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S. 6MIOIO 



^ GENERAL * 



Bulletin of the [; 7 JZ™ 

British Museum (Natural History) 



Botany series Vol 9 1981 



British Museum (Natural History) 
London 1982 



Dates of publication of the parts 

No 1 30 July 1981 

No 2 26 November 1981 

No 3 26 November 1981 

No 4 17 December 1981 



ISSN 0068-2292 



Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset 



Contents 
Botany Volume 9 

Page 
No 1 The lichenicolous Coelomycetes 

D. L. Hawksworth ......... 1 

No 2 The genus Callithamnion (Rhodophyta : Ceramiaceae) in the British 
Isles 
P. S. Dixon & J. H. Price 99 

No 3 Parmelia subgenus Amphigymnia (lichens) in East Africa 

H. Krog & T. D. V. Swinscow 143 

No 4 The genus Selaginella in tropical South America 

A. H. G. Alston, A. C. Jermy & J. M. Rankin .... 233 



Bulletin of the 

British Museum (Natural History) 



»0t 

GENERAL 

4 AUG mi 



The lichenicolous Coelomycetes 



D. L. Hawksworth 



Botany series Vol 9 No 1 30 July 1981 



The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
ever-growing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
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World List abbreviation : Bull. Br. Mus. nat. Hist. (Bot.) 



© Trustees of the British Museum (Natural History), 1981 



The Botany Series is edited in the Museum's Department of Botany 
Keeper of Botany: Mr J. F. M. Cannon 

Editor of Bulletin : Mr P. W. James 

Assistant Editor: Mr J. R. Laundon 



ISSN 0068-2292 Botany series 

Vol 9 No 1 pp 1-98 
British Museum (Natural History) 
Cromwell Road 
London SVV7 5BD I ssued 30 July 1981 



The lichenicolous Coelomycetes 

D. L. Hawksworth 

Commonwealth Mycological Institute, Ferry Lane, Kew, Surrey TW9 3AF* 



«> GENERAL 1 



Contents 

Synopsis I 

Introduction 2 

Biological and taxonomic concepts 2 

Methods 5 

Key to the genera 6 

Accepted species 7 

Ascochyta 7 

Asterophoma 8 

Bachmanniomyces 10 

Cornutispora 14 

Epicladonia 15 

Karsteniomyces 22 

Keissleriomyces 25 

Laeviomyces 26 

Libertiella 30 

Lichenoconium 33 

Lichenodiplis 37 

Lichenosticta 38 

Microcalicium [anamorphs] 41 

Minutophoma 44 

Nigropuncta 46 

Phoma 49 

Pseudoseptoria 57 

Pyrenotrichum 59 

Vouauxiella 64 

Vouauxiomyces 67 

Additional reports of anamorphs 71 

Excluded species 74 

Acknowledgements 89 

References 89 

Host index 92 

Fungus index 95 



Synopsis 

A revision of all Coelomycetes so far described from lichens is presented. Twenty genera are accepted, 
of which 16 are exclusively lichenicolous; seven of these are described as new and an additional new 
genus is introduced for the lignicolous Levieuxia borealis P. Karsten. Forty-two species are treated, 
including seven new species and seven new combinations; two additional new combinations are made 
for excluded taxa. A summary of the information on the anamorphs of 21 lichenicolous ascomycetes 
not considered in detail is included. Fifty epithets, including the type species often genera, are excluded 

*Present address: Commonwealth Agricultural Bureaux, Farnham House, Farnham Royal, Slough SL2 3BN. 



Bull. Br. Mus. nat. Hist. (Bot.) 9 ( 1 ) : 1-98 



Issued 30 July 1981 



2 D. L. HAWKSWORTH 

for a variety of reasons. Keys to the accepted genera and species are provided, as are indexes to both the 
lichen hosts and fungi reported as growing on them. 



Introduction 

Remarkable progress has been made in the last three years in synthesizing the available 
information on the Coelomycetes, fungi producing conidia within flask-like (pyenidia) or 
disc-like (acervuli) sporocarps (conidiomata). Sutton (1977) assessed the status of the 1336 
generic names proposed for Coelomycetes up to that time and was able to reject 720 for a 
variety of reasons. Synoptic keys to 200 genera were provided by Michaelides et al. (1979), 
and Sutton (1980) has prepared an exhaustive treatment of 375 accepted genera, including 
numerous line drawings and keys. Despite uncertainty in the application of 226 generic 
names, it is now possible to approach the Coelomycetes from a firmer taxonomic base than 
has hitherto been possible. 

It is against this background that the present revision of the Coelomycetes now known 
from lichens is presented. As in the case of my companion treatment of the lichenicolous 
Hyphomycetes (Hawksworth, 1979), this work is not to be interpreted as a definitive 
monograph. Rather, it should be viewed as a summation and re-assessment of the available 
information to provide a starting point for future work on this group of lichenicolous fungi. 

Vouaux (1914: 191-198, 281-301) treated 16 genera of lichenicolous Coelomycetes, 
comprising 52 species, world-wide. Keissler (1930: 534-589) accepted 42 species in central 
Europe distributed through 19 genera. In the compilation of Clauzade & Roux (1976: 
77-89). 54 species were distributed through 19 genera, although it should be stressed that 
their publication overlooked some previously published generic and specific names. Even at 
this late date, arbitrary nineteenth century generic concepts, ignoring methods of 
conidiogenesis and critical study of conidiomata, were still being employed. Furthermore, 
the identities of all but a handful of taxa had never been illustrated nor re-examined since 
their original description. 

In the course of the last few years, I have investigated a number of lichenicolous 
Coelomycetes (Hawksworth, 1975a, 1976, 1977, 1978a, 1980a; Hawksworth & Dyko, 
1979: Hawksworth & Punithalingam, 1973). The present work represents a culmination of 
my studies on these fungi by considering all names previously described as lichenicolous 
Coelomycetes and, further, by taking the opportunity to describe some hitherto 
unrecognized genera and species. 



Biological and taxonomic concepts 

Biological concepts 

Very few, if any, lichens appear to have anamorphs (imperfect states) referrable to the 
Hyphomycetes (Vobis & Hawksworth, 1981), so the separation of the normal anamorphs of 
the host from Hyphomycetes also present does not create any special problems. In the case of 
the Coelomycetes the situation is very different. Pycnidial anamorphs are known in perhaps 
as many as 8,000 lichens and these have repeatedly been misinterpreted as lichenicolous 
fungi: six generic and 19 species names are excluded here as definitely or probably based on 
the normal pyenidia of the host (pp. 74-89). 

There is a considerable degree of variation between the coelomycete anamorphs of lichens 
depending on the order to which they belong (Vobis & Hawksworth, 1981; Vobis, 1980). 
However, these structures have been little studied by lichenologists and only in the last few 
years have the) started to be described more fully in species descriptions. Fortunately there 
a few early papers which include careful drawings of a wide range of lichen pyenidia 
(Tulasne, 1852; Lindsay, 1859, 1872;Gliick, 1899;Gall0e, 1927-1972); it is essential when 



LICHENICOLOUS COELOMYCETES 3 

working on lichenicolous Coelomycetes to have these works to hand, and also to check 
herbarium specimens in the case of undocumented taxa. This is especially important when 
the invading fungus does not cause any visible damage to the host (i.e. is parasymbiotic). If 
necrotic patches, gall-like growths, or discolourations occur, or the pycnidia are within the 
hymenium or discrete local patches, it is fortunately normally safe to assume that a 
lichenicolous fungus is involved. 

In some cases the normal pycnidia may function as spermatia, but at least in some groups 
the conidia are evidently able to germinate to form extensive mycelia (Vobis, 1977). For 
descriptive purposes it is, following usage in other ascomycetes, now usual to use the terms 
'pycnidia' and 'conidia' as morphological ones not implying function. There seems to be 
no reason to depart from this procedure when considering lichen-forming fungi. 

Some lichens regularly occur in the pycnidial state (e.g. Cliostomum ghffithii (Sm.) 
Coppins) and there are examples where only pycnidia and no ascocarps are known (e.g. 
Lecanactis subabietina Coppins & P. James; Coppins & James, 1979); in the latter case the 
species are referred to genera defined on the ascosporic state only on the basis of similarities 
in the pycnidia and chemical products. Some knowledge of these cases is prerequisite to 
investigations into the lichenicolous Coelomycetes if unfortunate errors are not to be 
introduced. The situation is further complicated by the recent report of a probably dioecious 
lichen, Lecidea verruca Poelt (Poelt, 1980), and the probable existence of some true 
lichen-forming Coelomycetes. Many genera of the latter have been described, particularly 
from South America, but most merit re-investigation (Vobis & Hawksworth, 1981); one is 
treated as a synonym of Vouauxiella here (p. 64) and it is possible that some others also 
really represent lichenicolous fungi. One taxon described in the present revision, 
Nigropuncta rugulosa (p. 46), might conceivably be a lichen-forming coelomycete. 

Additional complexities arise from the occurrence of pycnidial anamorphs in 
lichenicolous ascomycetes. These have so far been described for very few such fungi, and 
usually very inadequately. Many more are to be expected to be recognized as the 
lichenicolous ascomycetes become better known. In the course of this revision I have not 
made a critical search for these and describe in full only a few which can be found without 
the teleomorph (ascosporic phase). The results of a preliminary literature survey are, 
however, included for completeness (pp. 71-74) as some of these reputed anamorphs can 
be expected to occasionally occur alone. 

The above considerations will make it clear that the production of a definitive account of 
obligately lichenicolous Coelomycetes will probably not be possible until (1) much more is 
known of the anamorphs of lichen-forming ascomycetes, (2) those of lichenicolous 
ascomycetes, and (3) an unequivocable basis for separating lichen-forming Coelomycetes 
from parasymbiotic lichenicolous taxa on 'unknown' thalli emerges. I doubt that the latter 
will ever be achieved in view of the complexity of fungus-alga interrelationships 
(Hawksworth, 1978/?). Fortunately, as stated at the start of this section, it is only where the 
invading fungus does not cause necrotic patches, discolourations, or gall-like growths that 
such biological demarcation problems arise. 

The lichenicolous Coelomycetes causing damage to the host can clearly be interpreted as 
parasitic (e.g. Lichenoconium erodens), but when the host is little affected with no 
discolouration or damage to the algal cells the situation is better referred to as parasymbiotic. 
In some instances (e.g. Pyrenotrichum splitgerberi) the invading fungus may depress 
ascocarp production in the host, have evenly scattered conidiomata, and appear lichenized 
itself; such instances may perhaps be compared to the production of one lichen by the 
usurping of the algae from a pre-existing lichen colony (Hawksworth et al., 1 980). 

Gall formation, especially well marked in the case of some lichenicolous Coelomycetes on 
Cladonia (e.g. Bachmanniomyces uncialicola, Epicladonia sandstedei), may be viewed as an 
example of parasymbiosis in which the fungus has an important role in morphogenesis. Galls 
appear to be particularly common in Cladonia where they can be induced by a wide range of 
fungi and perhaps also other organisms. Painstaking surveys of Cladonia galls were prepared 
by Bachmann (1927a, 19276, 1928a, 19286, 1929), but they occur more rarely in many 



4 D. L. HAWKSWORTH 

other genera also (Grummann, 1960). The causal agents of many lichen galls currently 
remains obscure; as knowledge of the lichenicolous fungi increases more may be related to 
individual species, but in some cases it may be a host reaction to invasion by disparate taxa 
which is being observed (cf. Hawksworth, 1980/?: 174). 

In m\ revision of the obligately lichenicolous Hyphomycetes (Hawksworth, 1979: 186), 18 
species were excluded as saprophytes found on dead or decaying thalli; several of these 
were ubiquitous fungi known from an extremely wide range of substrates. In the case of the 
Coelomycetes found on lichens this does not at present appear to produce any difficulties. As 
far as I am aware no coelomycete species also known from other substrates has yet been 
found on cither healthy or decaying lichen thalli. 

Taxonomic concepts 

The characters of value for the delimitation of genera in Coelomycetes have already been 
discussed in detail by Sutton (1973, 1980), whose works should be consulted for full 
information on this aspect. It is, however, necessary to draw attention to two particular areas 
of difficulty. During the last 1 5 years there has been an increasing emphasis on the method of 
conidiogenesis in the delimitation of genera of Coelomycetes. However, while the increased 
precision in the description of taxa is welcome and long overdue, in practice conidiogenesis 
in Coelomycetes is often much more difficult to study than in many Hyphomycetes as the 
conidiogenous cells tend to be very much smaller. When working close to the limits of 
resolution of light microscopy, it is all too easy to be misled; ultrastructural studies have 
demonstrated this in several cases. It is further becoming clear that some differences which 
have been given considerable taxonomic weight are not as clearly demarcated at the ultra- 
structural level as was at first presumed; the outstanding example in this case is the annellide/ 
phialide separation (Sutton, 1980: 20; Vobis & Hawksworth, 1981). Similar difficulties in 
the Hyphomycetes led Malloch (1979) and Kendrick (1980) to assert that too much 
emphasis should not be accorded to minor differences in conidiogenesis in otherwise very 
similar taxa. In the case of the Coelomycetes such caution is also required and I have 
endeavoured to bear this in mind in the present revision, as in my earlier account of 
Lichenoconium where the problem was also recognized (Hawksworth, 1977: 161). 

Sutton (1980) endeavoured to separate pycnidia-like conidiomata into two main types: 
simple (unilocular) eustroma when composed of a mass of fungal cells or closely interwoven 
hyphae, reserving 'pyenidia' for ostiolate, brown, thin-walled structures with walls 
composed of angular pseudoparenchymatous cells (textura angularis). Wall structure is an 
extremely important, but until recently largely overlooked, character in the Coelomycetes as 
in many Pyrenomycetes and Loculoascomycetes. Particular attention has consequently been 
accorded to this feature in the present study. The introduction of increasing numbers of 
terms for superficially similar but ontogenetically or structurally distinct sporocarps 
in the ascomycetes is now starting to be rejected, and it therefore seems unwise for 
coelomycete terminology to pass a similar course. In this account, c pycnidium' is 
consequently retained for all globose or flask shaped conidiomata regardless of the wall 
structure: the nature of the walls is of course indicated in the generic and species 
descriptions. 

Conidial shape, colour, septation, ornamentation, and whether extruded in mucous or 
dry, are all important characters which, taken in conjunction with others, support generic 
distinctions. It is, however, becoming clear that the abilities of conidia to develop additional 
septa or become slightly pigmented do not provide a sound basis for generic separations on 

eir own. In general, such features are rather of specific importance and these, taken 
together with their size and any differences in pyenidial sizes or aggregation, provide the 
species concept in most genera treated here. 

The ability of a fungus to occur on a particular host does not provide a specific criterion if 
other features are similar. For example, Cornutispora lichenicola was originally described 

nn Parmelia sulcata but is now also known from Lobaria pulmonaria, P. borreri, P. 
glabratula subsp. fuliginosa, Platismatia glauca, and Rhizoplaca chrysoleuca (see p. 14); 



LICHENICOLOUS COELOMYCETES 5 

Vouauxiomyces truncatus was thought to be restricted to P. caperata but has since been 
found on P. perlata (see p. 71). Where occurrences on different hosts are related to 
differences in conidial or pycnidial size or other features they can of course support species 
separations, as in the distinction of V. santessonii from V. truncatus. 

Reaction of the host to infection was recognized by Santesson (1949) as an important 
supporting feature at the species level in the taxonomy of lichenicolous fungi. This has 
already proved of value in Lichenoconium where species differing in pycnidial, conidio- 
genous cell, or conidial sizes give rise to dissimilar symptoms on the same host lichen 
(Hawksworth, 1977; Christiansen, 1980). 

It is with some hesitation that I introduce eight new generic names in this work, especially 
as I have already described three additional genera of lichenicolous Coelomycetes and 
resurrect two (Libertiella and Lichenosticta) not treated in recent reviews of coelomycete 
genera. However, this was to have been expected as (1) most lichenicolous fungi in other 
groups belong to obligately lichenicolous genera, and (2) the lichenicolous Coelomycetes 
have been largely ignored by earlier students of both lichenicolous fungi and other 
Coelomycetes. Additional obligately lichenicolous coelomycete genera certainly remain to 
be collected and described. Indeed, I have several unnamed specimens which probably 
represent new genera but are too scant to permit their thorough examination and 
description. At the same time it is possible that 'intermediate' species may be discovered 
necessitating the union of some genera accepted here (e.g. Laeviomyces and Lichenodiplis) 
and others may merit transfer to independent genera (e.g. Pseudoseptoria usneae, 
Vouauxiella uniseptatd). 

In the case of discovery of a lichenicolous coelomycete which does not appear in this 
revision, it must be stressed that before deciding it is undescribed it is of paramount 
importance to establish (a) that it is not the normal pycnidia of the host, and (b) examine the 
literature on other Coelomycetes (especially Sutton, 1973, 1980) to see if it can be 
accommodated into another not obligately lichenicolous genus (as is the case with e.g. 
Ascochyta lichenoides, Cornutispora lichenicola, Phoma species). 

Glossary 

Most terms which may be unfamiliar to lichenologists are defined in Hawksworth (1979: 
289-290); this glossary should be supplemented by that of Sutton (1980: 642-645) for 
additional terms peculiar to the Coelomycetes. 



Methods 

In order to ascertain the names of Coelomycetes described from lichens, the literature 
searches carried out for my survey of the lichenicolous Hyphomycetes were repeated (see 
Hawksworth, 1979: 189). Undoubtedly some taxa have been inadvertently overlooked, but I 
endeavoured to make this aspect of the work as thorough as possible so that the present 
revision can serve as a starting-point for further investigations on these fungi. 

Specimens supporting the various published reports were traced wherever possible and the 
material re-examined, supplemented by other collections available. Type or other material 
was obtained from the following herbaria in the course of this study: Angers, B, BM, E, GZU, 
H, IMI, K, LPS, M, PC, S, UPS, URM, SIENA, W, herb. Hafellner, herb. Poelt, and herb. 
Vouaux (the latter with Prof. Y. Rondon at Marseille). Herbarium abbreviations follow 
Holmgren & Keuken (1974). An exclamation mark (!) indicates that I have examined the 
specimen cited. 

The microscopic techniques used for the lichenicolous Hyphomycetes (see Hawksworth, 
1979: 189-190) were employed; the temporary erythrosin mountant was especially useful for 
the study of conidiogenous cells. Sections cut at lO/^rn with a freezing microtome were 
examined for almost all taxa discussed. During the latter part of this work I have been 
fortunate in being able to use an Olympus BH microscope fitted with Nomarski differential 



6 D. L. HAWKSWORTH 

interference contrast, drawing tube, and camera. The need for good optical equipment in 
studies on Coelomycetes cannot be over-emphasised. 

The scanning electron microscope has already proved a valuable aid in the study of those 
lichenicolous Coelomycetes with ornamented conidia (Hawksworth, 1977; Hawksworth & 
Dyko, 1 979), but no new scanning electron micrographs are included here. 

Key to the genera 

This is an entirely artificial key to the genera accepted in this revision. It does not include 
anamorphs of lichenicolous ascomycetes which normally occur in association with the 
teleomorph (perfect state), information on which is summarized separately (pp. 71-74). In 
the case of genera which are not obligately lichenicolous, the key is designed only to allow for 
the known lichenicolous taxa within them. 

1 Conidiomata pycnidial 2 

Conidiomata acervular or superficial and helmet-shaped 21 

2(1) Conidia shades of brown 3 

Conidia hyaline 7 

3(2) Conidia arising singly 4 

Conidia arising in long chains 6 

4(3) Conidia unicellular 5 

Conidia 1 -septate Lichenodiplis (p. 37) 

5(4) Conidia verruculose or echinulate; conidiogenous cells ± hyaline; pycnidial walls dark 

brown Lichenoconium (p. 33) 

Conidia smooth-walled; conidiogenous cells brownish; pycnidial walls subhyaline 

Laeviomyces (p. 26) 

6(3) Conidia unicellular and smooth-walled, or 1 -septate and rough-walled Vouauxiella (p. 64) 

Conidia multicellular, muriform, rough-walled Nigropuncta (p. 46) 

7(2) Conidia unbranched 8 

Conidia with a distinct stem and two apical arms Cornutispora (p. 14) 

8(7) Conidia septate 9 

Conidia unicellular 12 

9(8) Conidia 1 -septate 10 

Conidia 3-septate; conidiogenous cells subglobose to short-ampulliform, polyphialidic 

Keissleriomyces (p. 25) 

10(9) Pycnidial wall hyphal or composed of sclerenchymatous cells 11 

Pycnidial wall pseudoparenchymatous; conidiogenous cells phialidic, 4-8 x 2*5-3 //m; 
conidia rounded at both ends, narrowly ellipsoid, 8*5-10 x 3-4 um 

Ascochyta lichenoides (p. 8) 
11(10) Pycnidia superficial, shades of orange, with sclerenchymatous walls; conidiophores 

elongate and branched Karsteniomyces (p. 22) 

Pycnidia immersed to erumpent, brown to black, with hyphal walls; conidiophores 
short, usually absent Epicladonia (p. 15) 

(8) Conidiogenous cells arising singly, conidia acrogenous (borne only at the apex) . 13 

( onidiogenous cells arising in irregularly branched chains, conidia acro-pleurogenous 
(borne both apically and laterally); conidia lacriform . . Lichenosticta (p. 38) 

12) Conidia bacilliform, ellipsoid or obpyriform, rounded or truncated at the base . . 14 

( onidia lens-shaped to pyriform, pointed at the ends, adhering in mucilage 

Bachmanniomyces (p. 10) 



LICHENICOLOUS COELOMYCETES 7 

14(13) Pycnidia brown or black, superficial or immersed 15 

Pycnidia whitish to pale orange, with a deep reddish-brown ring around the ostiole, 
superficial Libertiella (p. 30) 

15(14) Conidia distinctly truncated at the base; conidiogenous cells holoblastic .... 16 

- Conidia with a rounded base; conidiogenous cells enteroblastic (phialidic) ... 18 

16(15) Conidia elongate-ellipsoid or obpyriform 17 

- Conidia cymbiform Pseudoseptoria (p. 57) 

17(16) Conidia obpyriform; pycnidial wall pseudoparenchymatous . . Vouauxiomyces (p. 67) 
Conidia elongate-ellipsoid; pycnidial wall hyphal Epicladonia (p. 15) 

18(15) Pycnidial wall entire, cells around the ostiole not separating 19 

- Pycnidial base formed by the conidiogenous cells alone, cells around the ostiole readily 

separating Minutophoma (p. 44) 

19(18) Conidiogenous cells subcylindrical or elongate-ampulliform 20 

Conidiogenous cells subglobose to short-ampulliform Phoma (p. 49) 

20(19) Pycnidial wall ± smooth; conidia subglobose or narrowly ellipsoid 

Microcalicium [anamorphs] (p. 41) 
Pycnidial wall with pointed cells projecting from it; conidia bacilliform 

Asterophoma (p. 8) 

21(1) Conidiomata helmet-shaped, superficial; conidia simple, multiseptate, or to 4-branched 

Pyrenotrichum (p. 59) 

Conidiomata scarcely developed, acervular; conidia with c. 50 radiating multiseptate 
arms Psammina (see Hawksworth, 1979: 244-246) 

Accepted species 

The accepted genera and species have been arranged alphabetically for ease of reference. In 
the case of genera including more than a single obligately lichenicolous species, keys to the 
species are included after the account of the genus itself. Descriptions and full synonymies 
for taxa I have previously treated elsewhere are generally omitted, but references to full 
descriptions, illustrations, etc., are provided. Synonyms not repeated from these works have 
been included in the Fungus index (pp. 95-98). Where genera include both lichenicolous 
and non-lichenicolous species, the generic diagnoses cover the genus as a whole and are 
partly based on those provided by Sutton (1980). 

I. ASCOCHYTA Lib 

PL Crypt. Ard.Jasc. 1 : 8 (1830). 

See Sutton (1980 : 408) for generic nomenclature. 

Conidiomata pycnidial, arising singly or in small groups, globose, immersed, brown to 
dark brown, ostiolate; walls composed of a few layers (often 1-3) of cells, cells pseudo- 
parenchymatous, pale to dark brown, rather thin-walled, textura angularis. Conidiophores 
absent. Conidiogneous cells enteroblastic, lining the pycnidial cavity, acrogenous, short- 
cylindrical to doliiform, phialidic, not proliferating, hyaline. Conidia arising singly, not 
catenate, narrowly ellipsoid or subcylindrical, rounded at the apex but often with a some- 
what truncated base, hyaline, 1 -septate when mature (exceptionally becoming 2-septatc). 
sometimes distinctly guttulate, thin-walled, smooth-walled. 

Type species: Ascochyta pisi Lib. 

Number of species: About 350 species are currently accepted but there are many more 
names in the literature. Taxa in the genus have generally been separated on a host basis in 
the past, and thorough revisions are required on the lines of that of Punithalingam (1979) 



8 D. L. HAWKSWORTH 

concerning the graminicolous species. Most species occur as leaf and stem-spotting fungi; 
only a single lichenicolous species is referred to this genus here. 

Observations: For practical purposes this genus may be viewed as a 1 -septate counterpart 
of Phoma (see p. 49), which it resembles closely in all other respects. As noted by Sutton 
(loc. cit.), attempts to separate these two genera on the basis of ultrastructural differences in 
conidiogenesis are difficult to relate to light-microscopical observations. 

1. Ascochyta lichenodes (A. L. Sm.) D. Hawksw., Notes R. bot. Gdn Edinb. 36 : 182 (1978). 

For illustration and further discussion of this species see Hawksworth {loc. cit.). 

Host: Indeterminate sterile crustose lichen on bark. 

Distribution: British Isles (England). Known only from the original collection. 

II. ASTEROPHOMA D. Hawksw. gen. nov. 

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata, 
dispersa vel laxe aggregata, globosa, superficialia vel hemi-superficialia, atrobrunnea vel nigra, 
ostiolata, cum muris atrobrunneis e cellulis elongatis pseudoparenchymaticis externe attenuatis et 
erumpescentis. Conidiophora desunt. Cellulae conidiogenae enteroblasticae, acrogenae, anguste 
ampulliformes vel late subcylindrices, phialidicae, interdum biphialidicae, non prolifericae, hyalinae. 
Conidia subcylindrica, hyalina, simplicia, laevia, minutissima, in muco aggregata. 

Conidiomata pycnidial, arising singly, uniloculate, scattered or loosely aggregated, 
globose, superficial or half-immersed, dark brown to black, ostiolate; walls composed of 
radially orientated elongated dark brown pseudoparenchymatous cells becoming attenuated 
and projecting slightly at the exterior to give the whole a star-like appearance. Conidiophores 
absent. Conidiogenous cells enteroblastic, interdigitated with the inner wall cells and lining 
the pycnidial cavity, acrogenous, narrowly ampulliform to broadly subcylindrical, phialidic, 
occasionally biphialidic, not proliferating, hyaline. Conidia subcylindrical, hyaline, simple, 
smooth-walled, minute, aggregated into a slimy mass which is extruded through the ostiole 
as a whitish drop. 

Type species: Asterophoma mazaediicola D. Hawksw. (holotypus). 

Number of species: Monotypic. 

Observations: This new genus recalls Minutophoma (see p. 44) in the minute 
conidiomata and subcylindrical conidia, but differs in the entire conidiomatal wall, presence 
of an ostiole, the externally attenuated wall cells, the hyaline conidiogenous cells that are 
also occasionally biphialidic, and the conidia being extruded in a mucilaginous whitish drop. 
It does not appear to have any other close allies. 

1. Asterophoma mazaediicola D. Hawksw. sp. nov. 

(Fig. 1A-D) 

Fungus lichenicola. Conidiomata singularia, uniloculata, dispersa vel laxe aggregata, globosa, super- 
ficialia vel hemi-superficialia, atrobrunnea vel nigra, 30-42 /zm diam, ostiolata, cum muris textura 
angulari 7-10 //m latis e cellulis atrobrunneis elongatis et externe attenuatis et erumpescentis 
5-8 x 2^1 //m. Conidiophora desunt. Cellulae conidiogenae enteroblasticae, acrogenae, anguste 
ampulliformiae vel late subcylindrices, phialidicae, interdum biphialidicae, non prolifericae, hyalinae, 

7 x 2*5-3*5 //m. Conidia subcylindrica, hyalina, simplicia, laevia, 15— 3(-3 5) x 0*5— 1 5 jum, in muco 
aggregata. 

Typus: Italia, Mendelgebirge, Nordhange des Roen, inter Mendel-Pass et Marga di Romeno, in 
ma/acdiis (aliens trabincllac Ach., 24.x. 1976, J. Hafellner 1822 (herb. Hafellner— holotypus!). 

( onidiomata pycnidial, superficial or partly immersed, arising in the mazaedial ascospore 
mass of the host, arising singly but sometimes loosely aggregated and juxtaposed, to 12 in a 
single mazaedium, dark brown to black, 30^2 jum diam, ostiolate; wall 7-10 //m thick, 
composed of a layer of radially elongated dark brown pseudoparenchymatous cells becoming 



LICHENICOLOUS COELOMYCETES 




D 



lOjLim 



Fig. 1 Asterophoma mazaediicola (herb. Hafellner — holotype). A, Infected mazaedium with 
pycnidia extruding conidia in mucilaginous drops. B, Vertical section of pycnidium. C, 
Conidiogenous cells. D, Conidia. 

attenuated and projecting slightly at the exterior to give the whole a star-like appearance, 
5-8 x 2-4 //m, lined internally with smaller subhyaline to hyaline subglobose to polyhedral 
pseudoparenchymatous cells 1 -5-3*5 jum diam; mycelium ramifying through the mazae- 
dium, hyphae hyaline, uneven in diameter, septate, irregularly branched, mainly 
1 *5— 25 jum thick. Conidiogenous cells enteroblastic, interdigitated with the inner wall cells 
lining the pycnidial cavity, difficult to distinguish, acrogenous, narrowly ampulliform to 
broadly subcylindrical, rather variable in shape, phialidic, occasionally biphialidic with two 
conidiogenous loci per cell, not proliferating, hyaline, 5-7 x 2*5-3*5 /zm. Conidia 
abundantly produced, subcylindrical, rounded at the apices, hyaline, simple, smooth-walled, 
not distinctly guttulate, l*5-3(-3*5) x 0*5-1 '5 jum, aggregated into a slimy mass which is 
extruded through the ostiole as a whitish drop. 

Hosts: Calicium glaucellum Ach. and C trabinellum Ach., apparently confined to 
mazaedial ascospore masses. Especially frequent on C trabinellum. Evidently a mild 
pathogen as diseased fruits often appear stunted and bent: the amount of vulpinic acid 
pigment produced by C trabinellum also appears to be reduced in diseased specimens so that 
such collections can easily be misidentified as C glaucellum. The absence of records from 
other Calicium species suggests that the fungus has a very narrow host range; indeed C 
glaucellum and C trabinellum are extermely close to one another and conceivably 



10 D. L. HAWKSWORTH 

chemotypes of a single taxon (Tibell, 1975 : 106). Infected fruits are easily recognized by the 
verrucosa surface that derives from the pycnidia themselves and particularly by the extruded 
white mucilaginous drops containing the conidia. 

Distribution: Canada, France, Italy, Norway, Rwanda, Sweden and the U.S.A. Probably 
scattered throughout the range of its host species (i.e. circumboreal), to judge from material 
so far available, and locally abundant. 

Observations: A distinctive species easily recognized by its very specialized habitat. The 
pycnidia of Microcalicium subpedicellatum (p. 42) are larger (40-60 jum diam), with a 
totally different wall structure, longer phialides, broadly ellipsoid to subglobose conidia, and 
occur only on the thalli (not in the mazaedium); M. subpedicellatum is known from 
Calicium glaucellum (Tibell, 1978 : 244) but has not been reported from C. trabinellum. The 
pycnidia of these Calicium species, when present, only occur on the thallus, not the 
mazaedium, and produce larger conidia c. 4-5 x 1 5 /zm (fide L. Tibell, in litt.). 

Additional specimens: Canada: British Columbia: Glacier National Park, Highway 1, 7 km NE of 
Rogers Pass, on Calicium glaucellum, 6 September 1972, L. Tibell 4959 (UPS!); Wells Gray National 
Park, Helmcken Falls, on C glaucellum, 19 September 1972, L. Tibell 5185 (UPS!); Pemberton Valley, 
c. 20 miles N of Pemberton, on C glaucellum, 17 September 1972, L. Tibell 5 180 (UPS!). —France: 
Haute Savoie : Chamonix, 0*5 km S of Planpra, Plan des Chablettes, on C trabinellum, 27 August 

1970. L. Tibell 4375 (UPS!); 15 km SSE of Chamonix, on C trabinellum, 26 August 1970, L. Tibell 
4383 (UPS!). — Italy: Trentino-Alto Adige : Ortlegruppe, Martelltal, on C. trabinellum, 2 October 

1971, L. Tibell 4466 (UPS!). —Norway: Hedmark, Engerdal par., 3'5 km SW of Hovdroseter, on C 
trabinellum, 29 July 1973, L. Tibell 5548 (UPS!). —Rwanda: Chaine des Birunga, Versant S of Bisoke, 
on C trabinellum, 25 February 1972, J. Lambinon 72/RW/883 (LG n.v., fide Tibell, in litt). 
— Sweden: Asele Lappmark : Frederika par., Vargan, 1*8 km SW of Flakatrask, on C. trabinellum, 5 
August 1972, L. Tibell 46\2 (UPS!). Dalarna : Hamra par., Hamra National Park, on C trabinellum, 
20 July 1973, L. Tibell 5433 (UPS!), 5448 (UPS!); loc. cit., on C. glaucellum, 20 July 1973, L. Tibell 
5424 (UPS!). Harjedalen: Ujusnedal par., 6 km NNE of Ljusnedal, Ormberget, on C. trabinellum, 25 
July 1973, L. Tibell 5503 (UPS!). Jamtland: Fors par., 10 km NNW of Fors, on C. trabinellum, 2 
August 1977, L. Tibell 7262 (UPS!). Norbotten: Overkalix par., 39 km NW of Overkalix, mire between 
Svartberget and O. Naverberget, on C trabinellum, 14 July 1977, L. Tibell 6762 (UPS!). Lule 
Lappmark: Muddas National Park, 5 km E of Porjus, on C glaucellum, 21 July 1977, L. Tibell 6962 
(UPS!): loc. at., at the little stream N of Muddusfallet, on C. trabinellum, 21 July 1977, L. Tibell 7026 
(UPS!). Uppland: Vange par., Fiby urskog, on C glaucellum 22 March 1964, L. Tibell 2044a (UPS!). 
— U.S.A.: Michigan : Keweenaw Co., Isle Royale, Rasberry Island, outside Rock Harbor, on C. 
trabinellum, 24 August 1972, L. Tibell 4656 (UPS!). 

III. BACHMANNIOMYCES D. Hawskw. gen. nov. 

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata, 
dispersa. globosa vel subglobosa, immersa, hyalina sed ostiolis aurantiacis vel brunneis, cum muris 
textura intricatis. ostiolata. Conidiophora desunt. Cellulae conidiogenae holoblasticae, acrogenae, 
elongato-ampulliformes, prolifericae, annellatae, hyalinae. Conidia late fusiformia ad pyriformia, 
plerumque asymmetrica, hyalina, simplicia, laevia, in mucosa. 

Conidiomata pyenidial (eustromatic), arising singly, scattered, globose to subglobose, 
immersed, hyaline but orange to brownish at the ostiole; walls composed of moderately 
thick-walled compactly entwined hyphae forming a textura intricata, hyaline except near the 
ostiole. Conidiophores absent. Conidiogenous cells holoblastic, lining the inner wall of the 
pyenidial cavity, acrogenous, elongate-ampulliform, somewhat broader at the base and 
tapering above, proliferating, annellate, hyaline, unbranched, occasionally septate. Conidia 

n> and adhering together, not catenate, lens-shaped to pyriform, attenuated at the apices, 
base narrowly truncate at the point of secession, often asymmetric, hyaline, simple, 
smooth-walled. 

Type species: Bachmanniomyces uncialicola (Zopf) D. Hawksw. (syn. Phyllosticta 
uncialicola Zopf— holotypus). 



LICHENICOLOUS COELOMYCETES 



11 



I 





Fig. 2 Bachmanniomyces uncialicola (W 1929/2 — holotype of Sirococcus lichenicola). A, 
Gall with pycnidia close to the apex of a branch, note the thinner stem above the gall 
(x 1 25). B, Gall with pycnidia arising laterally, the branch similar in thickness above 
and below the gall (x 1 2*5). 



Number of species: Monotypic. 

Observations: Bachmanniomyces has no very close allies amongst the non-lichenicolous 
Coelomycetes. The conidiogenous cells in Discosporium Hohnel (type species D. populeum 
(Sacc.) B. Sutton) are rather similar but the pycnidia in that genus are dark brown and 
aggregated into stromata. Sirococcus Preuss (type species S. stobi Units Preuss) was probably 
selected by Keissler (in Bachmann, 1929) for this fungus as he interpreted the conidia as 
catenate, but in Sirococcus the pycnidia are carbonaceous, lack a distinct ostiole, and the 
conidiogenous cells are branched (see Sutton, 1980). 

This new genus is named in honour of Edwald Th. Bachmann (1850-1937) in recognition 
of his pioneering work on gall-like structures in lichens, including investigations of the type 
species of Bachmanniomyces (Bachmann, 1929). 

1. Bachmanniomyces uncialicola (Zopf) D. Hawksw. comb. nov. 
(Figs. 2A-B, 3A-E) 

Phyllosticta uncialicola Zopf, Hedwigia 35 : 324 (1896). 
Type: British Isles, Scotland, Perthshire, Dunkeld, summit of Birnam Hill, on Cladonia uncialis, 
September 1858, W. L. Lindsay (E — lectotype!). 
Phoma uncialicola (Zopf) Vouaux, Bull, trimest Soc. mycol Fr. 30 : 198 (1914). 
Sirococcus lichenicola Keissler, in Bachmann, Arch. Protistenk. 66 : 469 (1929). 

Type: Sweden, Asele Lappmark, Kirchspiel Vilhelmina. auf Cladonia amaurocraea, Sommer 
1928, G. NilssonfG. DegeliusJ(W 1929/2— holotype!). 



12 



D. L. HAWKSWORTH 

B 




lOum 

Fiu. 3 Bachmanniomyces uncialicola (W 1929/2 — holotype of Sirococcus lichenicola). A, 
Gall on geniculately deformed branch. B, Vertical section of pycnidium. C, Vertical 
section of pycnidial wall. D, Conidiogenous cells. E, Conidia. 

Exsiccatae*: Arnold, Lick. Exs. no. 1021a (BM!; sub Cladonia uncialis f. biuncialis). — Sandstede, 

Clad. Exs. no. 161 (BM!; sub C. uncialis), no. 162 (BM!, UPS!; sub C. uncialis).— Schaerer, Lich. 

If civ. no. 514 (not found on this number in BM but cited by Sandstede, 1931 : 192; sub C. 

siellaia (i. ceranoides f. adusta). — Zwackh, Lich. Exs. no. 695 (n.v., cited by Sandstede, loc. 

til.: 192; sub C. uncialis f. leprosa). 
(cones: Bachmann. Bcr. dtsch. hot. Gcs. 38 : 335 fig. 6 [?] (1921); Arch. Protistenk. 66 : 461 figs 1-2, 

462 fig. 3. 465 figs 4-1 1 (1929).— Keissler, Rabenh. Krypt.-FL 8 : 537 figs 100-101, 553 fig. 

106a (1930).— Lindsay, Trans. R. Soc. Edinb. 25 : pi. 7 figs 15-16 (1859). 

Conidiomata pycnidial, completely immersed in gall-like deformations of the host, the 
galls massive irregular convex swellings to 1-1 "5 mm diam making the branches on which 
they occur geniculate, galls concolorous with the thallus, pycnidia arising singly, not 
aggregated into stromata, evenly scattered over the galls, the ostiole depressed below the 
surface level of the gall, orange at first but becoming brownish with age, 50-70 //m wide 
with the opening mainly 40-50 //m wide, pycnidia globose to subglobose, mainly 

er(1930 : 540) cited Sandstede Clad. Exs. no. 106; that number is C. portentosa Dufour (BM!) and does not 
support this fungus. 



LICHENICOLOUS COELOMYCETES 1 3 

100-200 //m diam when mature; pycnidial wall translucent, hyaline, mainly 12-20//m 
thick, becoming slightly more swollen and orange to brownish near the ostiole, the pigmen- 
tation due to interstitial mainly extra-hyphal deposition, only the limiting membrane of the 
hyphae in the ostiolar region coloured, wall throughout formed of irregularly orientated 
entwined and compacted moderately thick-walled hyphae mainly 2-3 //m wide. 
Conidiogenous cells holoblastic, arising from the inner wall of the pycnidium, completely 
lining the pycnidial cavity, elongate-ampulliform, the bases often expanded and tapering 
markedly towards the apex, hyaline, smooth-walled, annellidic with to at least 4 
annellations, unbranched, simple or occasionally 1 -septate, 7-12 jum tall, mainly 3^4- jum 
wide at the base and tapering to 2-3 jum at the apex. Conidia abundantly produced, not 
catenate, slimy and adhering together in large groups in aqueous preparations, lens-shaped 
to pyriform, often asymmetrical, apices often somewhat attenuated, the basal one more 
markedly so and also abruptly truncate when about 1 jum thick, hyaline, simple, often with 
a large guttule, smooth-walled, (7-)8-10(-105) x 4-5 -5 (-6) jum. 

Hosts: Cladonia amaurocraea (Florke) Schaerer, C. terrae- novae Ahti, and C. uncialis (L.) 
Wigg., thalli. The infection leads to the formation of distinctive galls (see above; Figs 2, 3). 
The galls are of the same colour as the healthy thallus and healthy algal cells consistently 
occur in the close proximity of the pycnidia; this fungus consequently appears to form a 
stable relationship with the host and not to be a pathogen. Vouaux's (1914 : 198) mention of 
its occurrence on C. bellidiflora (Ach.) Schaerer is due to confusion with ' Abrothallus" 
moorei Lindsay, and his mention of it on C. stellaris Opiz may also be incorrect. Keissler's 
(1930 : 540) reports from C. ochrochlora Florke, "C. sihaticcC and C. surrecta Florke, and 
Sandstede's (1931 : 3 1 7) from C. turgida (Ehrh.) Vainio, require confirmation. 

Distribution: British Isles (Scotland), Canada, France, Germany, Italy, Sweden and 
Switzerland. 

Observations: A very detailed account of the galls produced by Bachmanniomyces 
uncialicola is given by Bachmann (1929 : 460^69) who did, however, describe the conidia 
as forming in chains. I was unable to see any evidence of this, though the conidia strongly 
adhere in irregular slimy masses which presumably swell when moist and ooze through the 
rather wide ostioles. If the conidia were catenate, it would be anticipated that the mature 
ones would have a minute truncate point of secession at both ends when in fact this only 
occurs at their bases. The first-formed conidium arises holoblastically and if a young 
pycnidium is studied no or few conidiogenous cells with annellations will be found. The 
annellations are not easy to see in older conidiogenous cells unless studied in erythrosin- 
ammonia or a similar mountant. 

Lindsay (1859 : pi. 7 figs 15-16) was evidently the first author to recognize this fungus. He 
originally considered it to be a pycnidial state of Lecidea cladoniaria Nyl. (syn. Nesolechia 
cladoniaria (Nyl.) Arnold) but later realized that this was not so (Lindsay, 1871: 28-29). He 
also discovered an ascomycete on this host, named \4brothallus" moorei Lindsay (Lindsay. 
1869a : 545), and described its conidia which were, however, much longer and narrower 
than those of Bachmanniomyces uncialicola. Vainio (1887:265) recognized that defor- 
mations of Cladonia uncialis named as var. leprosa Delise were caused by a lichenicolous 
fungus, and Harmand (1907 : 245) ascribed this to Phyllosticta uncialicola Zopf. 

Zopf (1896:324) gave an extremely short diagnosis of Phyllosticta uncialicola: 
'Anschwellung an der Podetien bewirkend. Conidien einzellig, farblos, ei- oder birnformig. 
8, 4 : 5' and did not cite any specimens. No material of this taxon could be located in B (B. 
Hein, in litt.) but the original description is so characteristic of the present species that there 
can be no doubt as to the taxon he intended to describe. However, as Zopf introduced the 
name in his compendium of lichenicolous fungi then known, it is most unlikely that he 
overlooked Lindsay's (1859) description and comments on this fungus: Zopf cites these 
works in the introduction to his list and it is probable that he introduced the name with 
Lindsay's then unnamed fungus in mind. Lindsay's material is consequently designated 



14 D. L. HAWKSWORTH 

lectotype for Zopfs name here, but the possibility that it was the holotype should not be 
discounted. 

Vouaux (1914 : 198) indicated fc sans doute' that Spilomium epicladonia H. Olivier was a 
synonym of the present species. However, as the conidia of Olivier's fungus were given as 
6-8 x 3-4 um it seems unlikely that this interpretation is correct. 

Additional specimens: Canada: British Columbia, Queen Charlotte Islands, Graham Island, central 
part between Boulton Lake and Watun River, 53°49'N, 132° 4' W, on Cladonia uncialis, 3 August 
1966. //. Sjors (H!). Newfoundland, St. Mary's District, 2 mi E of Peter's River, on C. terrae- novae, 7 
June 1956. T. Ahti 9327 (H!). — Germany: Oldenburg, Kaihausermoor in Moorboden, on C. uncialis, 
April 1918, H. Sandstede [Clad. Exs. no. 162.] (BM!, UPS!); he. cit., October 1917, H. Sandstede 
[Clad. Exs. no. 161.] (BM!). — Italy: Hone zwischen Glashutten und Volsbach, sudlich von Bayreuth, 
auf Sandboden des braunen Jura, on C. uncialis, 6 September 1884, F. G. C. Arnold [Lich. Exs. no. 
1 02 1 a.] (BM!). —Sweden: Uppland, Uppsala, Witulfsberg, on C. uncialis, 1852, 77?. M. Fries (UPS!). 

IV. CORNUTISPORA Piroz. 

Mycologia 65: 763 (1973). ' * 

Conidiomata pyenidial, arising singly, subglobose, almost translucent to pale brown, 
immersed to almost superficial, opening by an irregular disintegration pf the upper part of 
the pyenidial wall; walls composed of thin-walled' pseudoparenchymatous cells, textura 
angularis. Conidiophores arising from the inner wall- of the pyenidial cavity, irregularly 
branched, septate, smooth-walled, hyaline, conidia forming acropleurogenously. Conidio- 
genous cells polyblastic, integrated, indeterminate,- broader towards the apices, hyaline, with 
1-4 apical denticles. Conidia arising singly, triradiate with a subcylindrical stem, two apical 
branches diverging at an obtuse angle, and a distinctly narrowed foot-cell, hyaline, simple, 
smooth-walled. 

Type species: Cornutispora limaciformis Piroz. 

Number of species: Two; C. limaciformis, which occurs in the apothecia of Therrya 
Juckelli (Rehm) Kujala (Phacidiales), and a single lichenicolous species distinguished by its 
much shorter conidia. 

Observations: The conidia of this genus are especially characteristic and it appears to have 
no close relatives in the Coelomycetes. 

1. Cornutispora lichenicola D. Hawksw. & B. Sutton, in Hawksworth, Trans. Br. mycol. Soc. 
67:51 (1976). 
(Fig.4A-B) 

This fungus is described and illustrated in Hawksworth (1976:51-53), and further 
drawings are given by Sutton (1980 : 90), so it is not discussed in detail here. 

Hosts: Lobana pulmonaha (L.) Hoffm. (thallus), Parmelia borreri (Sm.) Turner, P. 
glabratula subsp. fuliginosa (Fr. ex Duby) Laundon (thallus), P. sulcata Taylor (thallus), 
Plat is mat ia zlauca (L.) Culb. & C. Culb. (thallus), and Rhizoplaca chrysoleuca (Sm.) Zopf 
(syn. Lecanora rubina (Vill.) Ach.) (apothecia). Commonly in mixed infections with other 
lichenicolous fungi. The host tissues are sometimes blackened, decolourized, or otherwise 
discoloured, but it is often not clear whether this is due to the Cornutispora or the other fungi 
present. 

Distribution: Austria, British Isles (England, Scotland), Italy, and Switzerland. 

Specimens (additional to Hawksworth, loc. cit): Austria: Burgenland: Bez. Oberwart, SW of Bad 

rnannsdorf, on Parmelia sulcata, 7 April 1980, J. Poelt (GZU!; with Lichenoconium usneae). 

— British Isles: England, S. Devon, Slapton, Slapton Ley Nature Reserve, Peasdish, on P. borreri, 28 

\ugust 1980, /;. /.. Hawksworth 5064 (IMI 251276!; with L. lecanorae)- S. Somerset, Porlock, Horner 

Combe, on Platismatia glauca, 21 September 1980, D. L. Hawksworth 5072 (IMI 251490!); S. 



LICHENICOLOUS COELOMYCETES 



15 






Fig. 4 Cornutispora lichenicola (IMI 186831 — holotype). A, Conidiogenous cells. B, Conidia. 

Reproduced from Hawksworth (1976 : 52). 

Somerset, Nettlecombe Park, on Parmelia sulcata, 20 September 1980, D. L. Hawkswvrth 5069 (IMI 
251487!). Scotland, Kintyre, c. 7 km SW of Crinan, on islet N of Carsaig Island, on Lobaria 
pulmonaria, 14 August 1977, N. Brandt (E!, IMI 224707!; with Endophragmiella hughesii). — Italy: 
Sudtirol, Otztaler Alpen, Vinschgau, on Rhizoplaca chrysoleuca, 18 October 1975, J. Hafellner (GZU!; 
with Lichenoconium lecanorae). — Switzerland: Wallis, Aletschwald ob Brig, on P. glabratula subsp. 
fuliginosa, 1 1 September 1968,7. Poelt (hb. Poelt 6359!; with L. usneae). 



V. EPICLADONIA D. Hawksw. gen. nov. 

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata, 
subglobosa, ad cupuliformia, atrobrunnea immersa ad erumpescentia, cum muris textura intricatis, 
subhyalinis sed atrobrunneis in parte exposito, ostiolata. Conidiophora plerumque desunt. Cellulae 
conidiogenae holoblasticae, acrogenae, subcylindrices ad elongato-ampulliformes, prolifericae, 
annellatae, hyalinae. Conidia subcylindrica ad anguste-ellipsoidea vel cymbiformia, hyalina. 0-1 
septata, laevia. 

Conidiomata pycnidial (eustromatic), arising singly, scattered, subglobose to cupuliform, 
immersed to erumpent, dark brown; walls composed of compacted entwined short-celled 
hyphae forming a textura intricata, subhyaline except in the exposed parts, ostiolate. 
Conidiophores usually absent. Conidiogenous cells holoblastic, lining the inner wall of the 
pycnidial cavity, acrogenous, subcylindrical to elongate-ampulliform, proliferating, 
annellate, usually unbranched, hyaline. Conidia arising singly, not catenate, subcylindrical 
to narrowly ellipsoid or cymbiform, hyaline, 0-1 septate, smooth-walled. 

Type species: Epicladonia sandstedei (ZopO D. Hawksw. (syn. Pi plod ma mndstedei 
Zopf— holotypus). 

Number of species: Three, all of which are only known from Cladonia species on which 
they commonly form galls. 



16 D. L. HAWKSWORTH 

Observations: Epicladonia is distinguished from Bachmanniomyces, which also forms 
galls on Cladonia species (p. 10), in that the exposed parts of the pycnidia are dark brown to 
black, never shades of orange. Furthermore, the conidia are never pyriform nor strongly 
adhering in mucilage, 1 -septate in two species, and have a much broader scar than is seen in 
B. uncialicola. In Pseudoseptoria usneae (p. 57), the walls of the pycnidia are of 
greenish-brown pseudoparenchymatous cells (textura angularis) and the conidiogenous cells 
tend to be relatively short and stout. 

The three species referred to this new genus here are separated primarily on the basis of 
conidial size and septation and further, for E. simplex, the degree of emergence and extent of 
consequent pigmentation of the pycnidial wall. 

It is conceivable that Verrucaster Tobler might provide an earlier name for this genus, but 
in the absence of further information on the type species (see p. 88), I feel it is not possible 
to take it up here with any degree of confidence. 

key to the species 

1 Pycnidia mainly less than 100//m diam, immersed, brown only immediately around the 

ostiole 2 

Pycnidia 100-175 jum diam, erumpent and becoming almost superficial, the exposed parts 
all brown; forming galls; conidia simple, subcylindrical, rounded at the apex, 8-5—1 1(-12) 
x2 5-3(-3 5)//m Epicladonia simplex (p. 19) 

2 Conidia predominantly 1 -septate, subcylindrical to narrowly ellipsoid, apex always 

rounded, (7*5— )9— 1 2(— 1 4) x (2*5-)3-4 jum; usually forming galls Epicladonia sandstedei (p. 16) 

Conidia predominantly simple, subcylindrical to almost cymbiform, apex often attenuated, 
75-1 1 x 3-35 /zm; not forming galls Epicladonia stenospora (p. 20) 

1. Epicladonia sandstedei (Zopf) D. Hawksw. comb. nov. 
(Figs. 5A-B,6A-E) 

Diplodina sandstedei Zopf, in Sandstede, Abh. naturw. Ver. Bremen 18 : 429 (1906). 

Type: Germany, Oldenburg, Westerstede, 'Erdwalle in der Waldung "Hassen" bei Keilburg', on 
Cladonia caespitieia (Pers.) Florke [?], H. Sandstede. 
? Diplodina claudelii Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 289 (19 14); as k claudeli\ 

Type: France, Vosges, on Cladonia sp., V. Claudel. 

Exsiccatae*: Krypt. Exs. Vindob. no. 1330/? (sub Diplodina sandstedei; B!, K!). [The fungus was not 
found on copies of no. 1 330[a] studied (B!, K!).] 

hones: Keissler, Rabenh. Krypt.-Fl. 8 : 18 fig. 11,19 fig. 13, 25 fig. 20, 570 figs. 109-1 1 1 (1930). 

Conidiomata pycnidial, normally immersed in galls; galls convex, verruciform, to c. 1 mm 
diam. often somewhat constricted at the base, 5— 10(— 1 5) pycnidia in each gall, galls 
concolorous with the host tissue, not developed in early stages of infection when mature 
pycnidia can arise directly on the podetia; pycnidia arising singly, not aggregated into 
stromata, evenly scattered over the galls, immersed, the ostiole and upper part of the 
p\ cnidium slightly erumpent, translucent brown to almost black when mature, subglobose at 
first but becoming gaping and almost cupuliform with age, 50-80(-125)//m diam; ostiole 
clearly delimited in younger pycnidia, depressed, circular, to c. 50 jum diam, centre 
appearing subhyaline due to the conidial mass, formed by a breakdown of the upper wall of 
the pyenidium, irregular in shape in older pycnidia; pycnidial walls ± subhyaline except in 
the vicinity of the ostiole, there olivaceous brown to dark brown, mainly 8-1 5 jum thick, to 
20 fim thick near the ostiole, wall throughout composed of densely interwoven moderately 
thick-walled hyphae, these short-celled and mainly 2- 5^' 5 jum wide, textura intricata but 



"Sandstede 1 1 931) lists 24 exsiccatae numbers from seven sets on which he found this fungus. These are excluded as 
thc> were issued for the host Cladonia and the fungus is probably absent from some copies of these numbers. 



LICHENICOLOUS COELOMYCETES 




Fig. 5A-B, Epicladonia sandstedei (IMI 240228); A, infected podetium with galls (x 125); 
B, vertical section of pycnidium (x 500). C, E. stenospora (Dobbeler 1827), infected 
squamule showing decolorization and pycnidia (x 20). 

appearing almost cellular in parts. Conidiophores absent or short-cylindrical, simple or 
sparsely branched at the base, hyaline, variable in length, to 25 jum tall, 3^//m wide. 
Conidiogenous cells holoblastic, arising directly from the pycnidial wall or integrated into 
the conidiophores, subcylindrical to elongate-ampulliform, tapering towards the apex, 
hyaline, smooth-walled, annellidic, with 0-3 annellations, mainly 10-20//m tall, 3-4 /^m 
wide at the base, mainly 1 -5-2-5 //m wide at the apex. Conidia abundantly produced, 
arising singly, not catenate, subcylindrical to narrowly ellipsoid, rounded at the apex, 
truncated at the base with a scar 1 5—2 //m wide (remnants of wall tissue sometimes evident 
at high magnifications), hyaline, (0-)l — septate when mature, 2-3 guttulate, thin-walled, 
smooth-walled, (75-)9-12(-14) x (2'5-)3-4 //m. 

Hosts: Cladonia species, squamules and podetia, usually forming irregularly lobed bullate 
galls which may be hollow inside. Probably parasymbiotic. I have not made an exhaustive 
study of Cladonia specimens in search of the fungus but according to Sandstede (1931) it has 
a very wide host range. Sandstede mentioned it on C. anomaca (Ach.) Ahti & P. James, C. 
carneola (Fr.) Fr., C. coniocraea auct., C. chlorophaea (Florke ex Sommerf.) Sprengel s. lat., 



D. L. HAWKSWORTH 







lOjim 

Fig. 6 Epicladonia sandstedei. A-C (IMI 240228); A, vertical section of pycnidial wall; B, 
conidiogenous cells; C, conidia. D (Krypt. Exs. Vindob. no. 1330/?, K), conidia. E-F 
( 1 893. Harmand, Angers); E, conidiogenous cell; F, conidia. 



C. comma (L.) Hoffm., C. cyanipes (Sommerf.) Vainio, C. deformis (L.) Hoffm. , C.fimbriata 
(L.) Fr. (incl. C. major (Hagen) Sandst.), C. grayi G. K. Merrill ex Sandst., C. ochrochlora 
Florke. C. plcurota (Florke) Schaerer, C. pyxidata (L.) Hoflrn., C. subulata (L.) Wigg., and C. 
turgida Ehrh. ex Hoffm. 

His report from C. caespitica is probably an error for C. glauca Florke (see below), and 
at least one on C. pyxidata probably refers to C. merochlorophaea Asah. (see below); some 



LICHENICOLOUS COELOMYCETES 19 

other reports may also be due to mis-identified hosts or lichenicolous fungi. However, in 
addition, the fungus occurs also on at least C. conoidea Ahti (see below). 

Distribution: Evidently widespread in Europe. Reliably reported from at least the British 
Isles (England), France, Germany, Sweden and Switzerland. 

Observations: This fungus is easily distinguished from the other two species referred to the 
genus here on the basis of the predominantly 1 -septate conidia. 

The original material studied by Zopf could not be located in B (B. Hein, in litt.) but 
specimens from the type locality were distributed as Krypt. Exs. Vindob. no. 1330[a] (on 
Cladonia glauca Florke*; B!, H!, K!). Specimens on C. cf. merochlorophaea Asah., collected 
by Sandstede, were also issued as no. 1330/?*. No copy of 1330[a] seen had good material of 
the fungus, but as this exsiccatum would be ideal for neotypification, and the possibility that 
another copy with it well-developed exists, no neotypification is proposed here. The fungus 
on no. 1330/?, however, conforms to the usual concept of Diplodina sandstedei and so there 
is little doubt as to the correct application of the epithet. 

Diplodina claudelii is not present amongst the remnants of Vouaux's herbarium (Rondon, 
1970), nor could it be found in Claudel and Harmand's material in Angers (M. Guerlesquin, 
in litt.). This name was based on a fungus on the squamules of an unidentified Cladonia with 
pycnidia 90-1 50 pm diam, branched conidiophores with terminal conidia, conidiogenous 
cells 14-20 x 3 jum and 1 -septate conidia truncated at the base and rounded at the apex 
which measured 10-13 x 3-3*5 jum. There is consequently little doubt that this fungus 
should be placed here. The absence of galls does not preclude this possibility as these are not 
formed in early stages of infection; pycnidia in galls and directly on podetia can occur in the 
same collection (e.g. IMI 240228). As 1 -septate conidia were reported, it is unlikely that 
Vouaux was dealing with Epicladonia stenospora but that possibility cannot be 
entirely excluded as 2-guttulate conidia may appear septate if not studied carefully. 

I have not seen conidia over 1 1*5 jum in E. sandstedei, but Zopf (loc. cit.) originally gave 
them as 9-14 x 3*5^ jum. 

Keissler (1930 : 571) placed Diplodina peltigerae as a synonym of D. sandstedei, perhaps 
because of the similar conidial dimensions, but in the absence of material of the former and 
the disparate hosts such a treatment cannot be accepted here (see p. 76). 

Specimens: British Isles: England, W. Sussex, Pagham Harbour Nature Reserve, on Cladonia 
conoidea on soil amongst shingle, 30 August 1972, B. J. Coppins (IMI 240228!). —France: Docelles, on 
C chlorophaea s. lat., 1893, J. Harmand (Angers!). — Germany: See under Exsiccatae. 

2. Epicladonia simplex D. Hawksw. sp. nov. 
(Figs. 7A-B, 8A-B) 

Similis Epicladonia stenospora (Harmand) D. Hawksw. sed ditTert in pycnidiis erumpescentibus vel 
superficialbus, 100-1 75 jum diam, et conidiis semper simpliciibus, 85—1 1(-1 2) x 2*5-3(-3*5) /im. 

Typus: Finlandia,? Lapponia, in Cladoniae bacilliformes (Nyl.) Dalla Torre & Sarnth., c. 1925, V. 
Kujala (H — holotypus!). 

Conidiomata pycnidial, usually on galls formed on squamules or podetia of the host, galls 
convex, tuberculate, to c. 1 mm diam; pycnidia immersed at first but erumpent and almost 
superficial at maturity, arising singly, not aggregated into stromata, scattered, dark brown, 
subglobose to cupuliform, 100-175 pm diam; ostiole clearly delimited, depressed, gaping to 
reveal the subhyaline conidia inside and so appearing whitish centrally; pycnidial wall 
subhyaline in the parts sunken into the galls, but brown to dark brown in the exposed parts, 
mainly 10-1 5 pm thick, composed of densely interwoven short-celled moderately thick- 
walled hyphae mainly 2*5-3*5 jum wide, textura intricata, but becoming almost cellular in 
parts. Conidiogenous cells holoblastic, arising from the pycnidial wall, subcylindrical 

*Det. T. Ahti; the original host and material in no. 1330[fl] was erroneously named as C caespiticia (Pers.) Florke, 
and in no. 1 330/? as C. pyxidata (L.) HofTm., by Sandstede. 




D. L. HAWKSWORTH 



B 




Fig. 7 Epicladonia simplex (H — holotype). A, Infected podetia showing pycnidia arising 
from galls and directly on the podetia (x 12*5). B, Vertical section of pycnidium (x 500). 



to elongate ampulliform, only slightly tapered towards the apex, annellate with to 3 
annellations, hyaline, smooth-walled, 7-15 x2'5-3*5 jum. Conidia arising singly, not 
catenate, subcylindrical, always rounded at the apex, truncated at the base with a scar c. 
1-5-2 pm wide, always simple, 2 or more guttulate, thin-walled, smooth-walled, 8*5- 

ll(-12)x2-5-3(-3-5)//m. 

Host: Cladonia bacilliformis, usually forming galls on both the primary squamules and 
podetia of the host. Probably parasymbiotic. 

Distribution: Finland. Known only from the original collection. 

Observations: This species is most similar to Epicladonia stenospora (see below) from 
which it is distinguished by the generally larger and erumpent to almost superficial pycnidia, 
the more extensively pigmented and more cellular appearance of the pycnidial wall, the 
shape of the conidia, and further in giving rise to the formation of galls, although these are 
not formed immediately (Fig. 7A). 

The annellations on the conidiogenous cells are more distinct in this species than in E. 
sandstedei and none were noted in E. stenospora. Indeed, one might interpret the 
conidiogenous cells of E. simplex as proliferating phialides rather than annellides. However, 
I am sceptical as to the merits of such characters at the genus level (see p. 4) and so place this 
species here. 

3. Kpicladonia stenospora (Harmand) D. Hawksw. comb. nov. 
(Figs. 5C,9A-B) 

Aposphaeria stenospora Harmand, Lick Fr. 3 : 308 (1907). 

Conidiomata pycnidial, immersed in squamules, not associated with galls, arising singly, 
not aggregated into stromata, scattered, the ostiole slightly erumpent or depressed, brown, 
subglobose, 75-1 20 /^m diam; ostiole clearly delimited at first but later tending to gape; 
pycnidial wall hyaline except in the vicinity of the ostiole where it is olivaceous brown to 
dark brown, mainly 7-1 5 pm thick but to 25 jim thick near the ostiole, wall mainly 
composed of densely interwoven hyphae mainly 2-3 //m wide, textura intricata but 
becoming almost cellular in the vicinity of the ostiole and there 4-6 jum diam. 
Conidiogenous cells holoblastic, arising from the pycnidial wall, elongate-ampulliform, 



LICHENICOLOUS COELOMYCETES 



21 




Fig. 8 Epicladonia simplex (H- 



■holotype). A, Vertical section of pycnidial wall. B, Conidiogenous 
cells. C, Conidia. 



strongly tapered towards the apex, hyaline, smooth-walled, no annellations seen, mainly 
7-12 x 3-3 -5 jum. Conidia arising singly, not catenate, subcylindrical to almost cymbiform, 
often rather irregular in shape, the apex rounded to slightly attenuated and the base 
narrowed and abruptly truncated by a scar c. 1-1*5 jum wide, simple or exceptionally 
1 -septate, frequently 2 or more guttulate, thin-walled, smooth-walled, 7*5-1 1 x 3-3*5 jum. 

Hosts: Cladonia cf. coniocraea auct. and C. pyxidata (L.) Hoffm. [?] (see below). Occurring 
on the primary squamules in the specimen from Austria examined; no gall-like deformations 
are produced, but the infected tissue eventually becomes decolourized to give circular pale 
brownish necrotic infection spots on the host squamules (Fig. 5C). Probably parasitic. 

Distribution: Austria and France. 

Observations: Despite the lack of annellations this fungus is referred to Epicladonia in 
view of the very similar structure of the pycnidia and the occasional presence of 1 -septate 
conidia. Further, the presence of distinct annellations in E. simplex which consistently has 
simple conidia and further forms galls on Cladonia adds weight to this taxonomy. The 
separation of E. stenospora from E. simplex is discussed under the latter species (p. 20). 

In the original description of Aposphaeria stenospora, the host was named as Cladonia 
pyxidata but no locality details were provided. The original material could not be located in 
Angers (M. Guerlesquin, in titt.), but as the conidia were described as 8-9 x 2-25 pm it is 
probable that the name referred to a fungus such as that of Dobbeler's (see below), on which 
the above description is based. The Austrian specimen is not designated as a neotype for 
Harmand's name as it is rather sparse. 

Specimen: Austria: Tirol, Hange unmittelbar ostlich des Achensees in der Nahe des Achenseehofes, 
on Cladonia cf. coniocraea, 1 3 October 1976, P. Dobbeler 1 827 (GZU!). 



22 



D. L. HAWKSWORTH 





lOjwn 

Fig. 9 Epicladonia stenospora (Dobbeler 1827). A, Vertical section of pycnidial wall with 

conidiogenous cells. B, Conidia. 



VI. KARSTENIOMYCES D. Hawksw. 

Trans. Br. mycol. Soc. 74 : 371 (1980). 

Conidiomata pycnidial, arising singly, scattered or loosely aggregated, subglobose, 
superficial, nectrioid, translucent pale orange to deep red, ostiole irregular and forming 
schizogenously; walls thick, composed of thick-walled pseudosclerenchymatous cells with 
much-reduced lumina. Conidiophores cylindrical, sympodially branched. Conidiogenous 
cells holoblastic, arising acrogenously or pleurogenously, cylindrical, hyaline. Conidia 
probably dry, elongate-ellipsoid, the apex rounded and the base slightly to clearly truncate, 
hyaline, 1 -septate, smooth-walled, guttulate. 

Type species: Karsteniomyces peltigerae (P. Karsten) D. Hawksw. (syn. Stagonopsis 
peltigerae P. Karsten). 

Number of species: Monotypic. 

Observations: This genus appears to occupy an isolated position in the Sphaeropsidales 
and is unlikely to be confused with any other. Stagonopsis Sacc. (type species S. pallida 
(Berk. & M. A. Curtis) Sacc.) is of somewhat uncertain application, but as the type species 
was described as forming arcuate 7-8 septate conidia, it is most unlikely that it has anything 
to do with Karsteniomyces. 

This new genus is named in honour of Petter A. Karsten (1834-1917) who made very 
substantial contributions to our knowledge of microfungi in Finland, including the discovery 
of several previously unrecognized lichenicolous species. 

1. Karsteniomyces peltigerae (P. Karsten) D. Hawksw., Trans. Br. mycol. Soc. 74:371 
(1980). 
(Figs. 10A-D, 11A-E) 

Stagonopsis peltigerae P. Karsten, Meddel Soc. Fauna Flora fenn. 14 : 106 (1887). 
Type. Finland. Tavastia australis, Tammela, Mustiala, on Peltigera rufescens, July 1868, H. A. 
A'h///k'w(H-KARST2041— holotype!). 

Conidiomata pycnidial, almost entirely superficial with only the base entering the host 
tissues, arising singly but often numerous and sometimes crowded together, globose, 
translucent pale orange at first, becoming a dark orange red with age especially around 



LICHENICOLOUS COELOMYCETES 





% Mm 



*3? 



t.-~~G''. 










Fig. 10 Karsteniomyces peltigerae (H-KARST 2041 — holotype). A, Pycnidia on the host 
thallus (x25). B, As A showing the wide ostioles (x 50). C, Vertical section of pycnidium 
(x 250). D, Vertical section of pycnidial wall showing the thick-walled almost sclerenchymatous 
cells (x 1020). 



the ostiole, mainly 75-200 //m diam but exceptionally larger, appearing nectriaceous, 
opening by an irregular schizogenously formed ostiole; pycnidial wall very variable in thick- 
ness, mainly 20-50 jum thick, composed of two layers, the outer comprising ± hyaline very 
thick-walled pseudosclerenchymatous cells with extremely narrow compressed lumina, the 
outer layer sometimes almost absent, very uneven externally, 5-20 pm thick, the inner 
layer also pseudosclerenchymatous but formed of cells with larger lumina and pigmented 
pale orange, 10-25 jum thick. Conidiophores arising from the inner walls of the pycnidial 
cavity, cylindrical, irregularly sympodially branched, hyaline, smooth-walled, 5-15 pm tall 
and 2-3(-4)//m wide. Conidiogenous cells arising terminally and laterally from the 
conidiophores, indistinguishable from the conidiophores except in the absence of septa. 
Conidia holoblastic, abundant, dry or at least not adhering, elongate-ellipsoid, the apex 
rounded and the base slightly to clearly truncate, hyaline, 1 -septate when mature, the 
septum thin and sometimes undulate, \^\ guttulate, smooth-walled, (12-)1 5-22(-24) x 
3-5-5(-6)/im. 



24 



D. L. HAWKSWORTH 




Fif>. 11 Karsteniomyces peltigerae (H-KARST 2041 — holotype). A, Vertical section of 
pycnidium. B, Cells from the inner part of the pycnidial wall. C, Cells from the outer 
part of the pycnidial wall. D, Conidiophores and conidiogenous cells. E, Conidia. 



Host: Peltigera canina (L.) Willd. and P. rufescens (Weis) Humb., thalli. The type 
specimen was originally stated to be on P. canina but that host has been redetermined by Mr 
O. Vmkainen as P. rufescens. The species is, however, correctly reported here from P. 
canina (sec Additional specimens). The infected thalli do not appear to be very significantly 
damaged by Karsteniomyces peltigerae and no clearly demarcated decolourized patches are 
produced. 



LICHENICOLOUS COELOMYCETES 25 

Distribution: Finland and Norway. Keissler (1930 : 585) mentions two dubious records 
from Germany and Italy but these are in need of confirmation. 

Observations: The conidia have been described as becoming up to 4-septate (Keissler, 
1930:585) but I have only seen 1 -septate conidia. I strongly suspect that reports of 
additional septa are due to misinterpretations of the guttules, which are often conspicuous. 

Additional specimens: Finland: Regio aboensis, Kakskerta Kokkila, on Peltigera canina and P. 
rufescens, 4 August 1934, L. E. Kari (H — two packets!). — Norway: Akershus, Oslo (Christiania), 
Disen, on P. canina, 25 March 1 869, N. G. Moe 145 (UPS!). 

VII. KEISSLERIOMYCES D. Hawksw. gen. nov. 

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidomata singularia, uniloculata, 
dispersa, subglobosa, immersa, nigra, ostiolata, cum muris tenuibus et textura angulari compressis. 
Conidiophora desunt. Cellulae conidiogenae enteroblasticae, subglobosae vel breve ampulliformes, 
polyphialidicae, hyalinae. Conidia probaliter sicca, late fusiformia, basi leviter arcuata, hyalina, 
3-septata, laevia. 

Conidiomata pycnidial arising singly, scattered, subglobose, immersed, black, ostiolate; 
walls thin, composed of radially compressed polyhedral dark brown cells, textura angularis. 
Conidiophores absent. Conidiogenous cells enteroblastic, subglobose or short-ampulliform, 
polyphialidic, each giving rise to up to 3 conidia, hyaline. Conidia probably dry, broadly 
fusiform, the base weakly curved, hyaline, 3-septate, smooth-walled. 

Type species: Keissleriomyces sandstedeanus (Keissler) D. Hawksw. (syn. Stagonospora 
sandstedeana Keissler — holotypus). 

Number of species: Monotypic. 

Observations: The genus Stagonospora (Sacc.) Sacc. nom. cons, (type species S. paludosa 
(Sacc. & Speg.) Sacc.) is characterized by conidiogenous cells forming conidia holo- 
blastically and not polyphialides; S. sandstedeana consequently does not belong there. 
Keissler's choice of Stagonospora was most probably due to the rather similar conidia seen 
in some species of that genus. Polyphialidic conidiogenous cells are rather rare in 
Coelomycetes and Keissleriomyces has little else in common with other genera exhibiting 
this feature. 

Although with hindsight many of Karl von Keissler's (1872-1965) taxonomic decisions 
amongst the lichenicolous fungi prove to be unsatisfactory, his compendium of the central 
European species (Keissler, 1930) remains an essential reference work to all students of 
lichenicolous fungi today; it is in recognition of this debt to him that the new generic name 
Keissleriomyces has been introduced here. 

1. Keissleriomyces sandstedeanus (Keissler) D. Hawksw. comb. nov. 
(Figs. 12, 13A-E) 

Stagonospora sandstedeana, Keissler, Annln naturhist. Mus. Wein 38 : 166 (1925). 
Type: Czechoslovakia (Bohemia), Riesengebirge, Schneekoppe, on Cladonia furcata, O. I.. .1. 
Ohlert(W 1921/185— holotype!). 

Conidiomata pycnidial, almost entirely immersed in the host tissues, arising singh. 
scattered, subglobose but often radially compressed, black, 40-75(-80)/*m diam, ostiolate; 
pycnidial wall 7-10 jum thick, composed of usually two layers of radially compressed 
polyhedral pseudoparenchymatous cells, cells dark brown, moderately thick-walled. 
10-14 x 3-4 jum in vertical section and mainly 6-9 jam diam in surface view. Conidio- 
genous cells enteroblastic, lining the inner wall of the pycnidial cavity except near the apex, 
subglobose or shortly ampulliform with swollen bases, 5-8 x 5-6 um, polyphialidic. each 
cell with 1-3 phialides, hyaline. Conidia abundant, probably dry. broadly fusiform with the 
base generally more attenuated and frequently weakly curved, hyaline, 3-septate when 



26 



D. L. HAWKSWORTH 



? * 




Fig. 12 Keissleriomyces sandstedeanus (W 1921/185 — holotype), vertical section of pycnidium 

(x 1020). 



mature, not markedly guttulate, contents rather granular, smooth-walled, 18-22 x 
3-5—5 //m. 

Host: Cladonia furcata (Huds.) Schrader, podetia. In the type material the pycnidia are 
mainly located on the upper part of the podetia, the infected part of which is slightly 
decolourized but not deformed. Keissleriomyces sandstedeanus consequently appears to be 
only a weak parasite or opportunist. 

Distribution: Czechoslovakia. Only known from the type collection. 

Observations: The characteristic polyphialides are otherwise unknown amongst the 
lichenicolous Coelomycetes, and no other fungus forming similar spores is recorded from 
any Cladonia species. 

VIII. LAEVIOMYCES D. Hawksw. gen. nov. 

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, pycnidiifor- 
mia. uniloculata, dispersa, subglobosa vel irregularia, erumpescentia, nigra, cum muris plusminusve 
hyalinis e cellulis pseudoparenchymaticis (sed in unica textura intricatis) et ostiolis irregulariter 
dchiscentibus. Conidiophora desunt. Cellulae conidiogenae holoblasticae, acrogenae, subcylindrices 
vel elongato-ampulliformes, prolifericae, annellatae, fuscae vel brunneae, laeves. Conidia subglobosa 
ad ellipsoidea, basi truncata, pallide brunnea, simplicia, laevia. 

Conidiomata pycnidial, arising singly, uniloculate, scattered, subglobose or irregular with 
somewhat convoluted internal walls, erumpent, black; walls composed of several layers of 
irregularly polyhedral to subglobose ± hyaline cells forming a textura angularis (but in one 
species reduced and hyphal, textura intricata); ostiole irregular in shape and gaping in age. 
Conidiophores absent. Conidiogenous cells holoblastic, lining the inner wall of the pycnidial 
cavity, acrogenous, subcylindrical to elongate-ampulliform, proliferating, annellate, fuscous 
brown to brown, smooth-walled. Conidia subglobose to ellipsoid, abruptly truncated at the 
base, a small basal frill sometimes discernible, not catenate, brown, simple, smooth-walled. 



LICHENICOLOUS COELOMYCETES 



27 




Fig. 13 Keissleriomyces sandstedeanus (W 1921/185 — holotype). A, Vertical section of 
pycnidium. B, Vertical section of pycnidium wall. C, Surface view of pycnidial wall. D, 
Conidiogenous cells. E, Conidia. 



Type species: Laeviomyces pertusariicola (Nyl.) D. Hawksw. (syn. Spilomium pertusarii- 
cola Nyl. — holotypus). 

Number of species: Two, both lichenicolous. 

Observations: The type species of this genus was retained in Lichenoconium as an interim 
measure by Hawksworth (1977) because a suitable generic name was considered to possibly 
exist amongst the non-lichenicolous Coelomycetes. Subsequent studies have failed to reveal 
any suitable generic name and so the genus Laeviomyces is introduced here to accommodate 
it. The generic name recalls the smooth-walled conidia (x 8000), but L. pertusariicola further 
differs from Lichenoconium in the massive pycnidia, hyaline and relatively flaccid pycnidial 
wall, the consistently brown and smooth-walled conidiogenous cells, and individually much 
less strongly pigmented conidia, sometimes with a recognizable basal frill. 

Farr et al. (1979 : 1649) listed Spilomium pertusariicola Nyl. as the type species of 
Spilomium Nyl. However, they give the place of publication of the latter as Nylander 
(1858 : 91) when in fact the genus must be considered as already validly published earlier by 
Nylander (1856 : 337), who made no mention of S. pertusariicola although referring three 
other species to Spilomium. Nylander's generic name is correctly interpreted as a 
superfluous name for the hyphomycete genus Sclerococcum Fr. ex Fr., as already pointed out 
by Hawksworth (1975a : 222). 

Laeviomyces could be viewed as a counterpart of Lichenodiplis with simple as opposed to 
1 -septate conidia. In view of the more complex wall structure of Laeviomyces pertusariicola. 



28 



D. L. HAWKSWORTH 




3^ ^^wPV-fc-j* ■ 









Fig. 14 Laeviomyces opegraphae (E — holotype), vertical section of Opegrapha niveoatra 
lirellum with the hymenium partly replaced by a pycnidium of the Laeviomyces (x 500). 

and the emphasis placed on spore septation in traditional coelomycete systematics, it seems 
preferrable to retain two genera while intermediate taxa are unknown. 

Key to the species 

! Pycnidia 40-60 //m wide; pycnidial wall poorly developed and hyphal; conidia olivaceous 

brown, 3-4-5 x 1 5—2 jum Laeviomyces opegraphae (p. 28) 

Pycnidia 100-250(-325)/mi diam; pycnidial wall well developed and pseudoparen- 
chymatous; conidia reddish brown, 3 - 5-6 x 2*5-3 - 5 jum . Laeviomyces pertusariicola (p. 29) 



1. Laeviomyces opegraphae D. Hawksw. sp. nov. 

(Figs. 14J5A-B) 

Similis Laeviomycei pertusariicola (Nyl.) D. Hawksw. sed differt in pycnidiis 40-60 jum latis, muris 
rudimentalibus e textura intricata, et conidiis olivaceo-brunneis, 3-4*5 x 1 5—2 pirn. 

Typus: Magna Britannica, Scotia, Dumbarton, Loch Lomond, Gartocharn, Claddochside, in thallo 
Opegraphae niveoatrae (Borrer) Laundon, 3.iii.l980, B. J. Coppins 467 5 (E — holotypus!). 

Conidiomata pycnidial, immersed in the thallus or lirellae of the host, arising singly, 
scattered, sometimes becoming confluent in heavy infections, brown, 40-60 jum wide, to 
100/^m tall when in lirellae, at first globose but becoming very irregular in shape, ostiole 
forming by an irregular breakdown of the upper wall of the pycnidium, becoming gaping and 
the pycnidia sometimes almost cupuliform or melanconiaceous with age; pycnidial wall 
poorly delimited, composed of 2-3 layers of intertwined short-celled hyphae forming a layer 
only 3-5 pm thick, hyphae olivaceous, sometimes becoming almost cellular just below the 
conidiogenous cells, subhyaline to pale olivaceous brown, L5-2'5//m thick. Conidio- 
genous cells holoblastic, acrogenous, subcylindrical, proliferating, distinctly annellate with 
to 6 annellations, olivaceous brown, smooth-walled, 4-7(-9) x F5-2-5 //m. Conidia dry, 
ellipsoid, narrowed to an abruptly truncated base, basal frill sometimes visible, olivaceous 
brown, simple, not conspicuously guttulate, smooth-walled, 3^*5 x L5-2 jum. 

Host; Opegrapha niveoatra (Borrer) Laundon, thallus and hymenium of the lirellae. When 
on the thallus little damage occurs, but when the lirellae become infected almost the whole 
of the hymenium may eventually be occupied by the fungus (Fig. 14), thus precluding the 
possibility of ascospore formation. 



LICHENICOLOUS COELOMYCETES 



29 





10/im 

Fig. 15A-B, Laeviomyces opegraphae (E — holotype); A, conidiogenous cells; B, conidia. 

C, L. pertusariicola (IMI 1 86240), conidia. 



Distribution: British Isles (Scotland). Evidently not uncommon in Scotland on this host 
and to be expected throughout the range of the host lichen. 

Observations: This new species is included within Laeviomyces with some hesitation as 
the pycnidial wall is so poorly developed and almost hyphal, rather than pseudoparen- 
chymatous as in L. pertusariicola. However, the pigmented and annellate conidiogenous 
cells and shape of the conidia are so similar in these two fungi that it seems more appropriate 
to view L. opegraphae as a species in which the wall has been reduced. It would not be 
prudent to describe a new genus based only on this single feature in our present state of 
knowledge of the lichenicolous Coelomycetes. 

Additional specimens: British Isles: Scotland, Midlothian, Dalkeith Oak Wood, on Opegrapha 
niveoatra on Acer, 25 February 1977, B. J. Coppins & C. Pope 4696 (E!); Peebles, Dawyck Arboretum, 
on O. cf. niveoatra on A. pseudoplatanus, 10 May 1980, B. J. Coppins 4860 (E!). 

2. Laeviomyces pertusariicola (Nyl.) D. Hawksw. comb. nov. 
(Fig. 15C) 

Spilomium pertusariicola Nyl., Mem. Soc. Imp. Sci. nat. Cherbourg 5 : 91 (1858); as "pertusaricola\ 
Lichenoconium pertusariicola (Nyl.) D. Hawksw., Trans. Br. mycol. Soc. 65 : 233 (1975). 

For description, fuller illustrations, and further synonyms see Hawksworth (1975a: 
233-234,1977: 182-183). 

Hosts: Buellia disciformis (Fr.) Mudd, Pertusaria leioplaca DC. and P. pertusa (Weigel) 
Tuck., thalli. Apparently parasymbiotic, not damaging the host thalli. 

Distribution: British Isles, Canada, Denmark, France, Germany and Spain. There is also 
an unconfirmed report from New Caledonia (Keissler, 1933 : 387). 

Observations: In the Canadian collection on Buellia disciformis, a host now confirmed for 
this fungus, the conidia tend to be slightly longer (to 7*5 jum) and rather more deeply 
pigmented than is usual for the species; further collections on that host may show that it 
merits recognition as a separate taxon. Muellerella lichenicola (Sommerf. ex Fr.) D. 
Hawksw. is also present in the collection on this Buellia and intimately mixed with the 
Laeviomyces. 

Lichenodiplis lecanorae, which can also occur on corticolous Pertusaria species, is easily 
separated from Laeviomyces pertusariicola by the consistently 1 -septate and generally paler 
rather more cylindrical and longer conidia, which have an even more strongly truncated 
base, as well as by the much smaller pycnidia. 

Specimens (additional to those listed by Hawksworth, 1975a, 1977): British Isles: Scotland, 
Dunbartonshire, Loch Lomond, Ross Priory, on Pertusaria leioplaca on Acer, 13 March 1980, B. J. 
Coppins 4664 (E, IMI 247022!); Wales, Breconshire, Erw Fawr, above Elan Valley, on P. cf. leioplaca 



30 D- L. HAWKSWORTH 

on Quercus, 20 October 1979, F. Rose (IMI 244540!). —Canada: Nova Scotia, Halifax, South Park, on 
Buellia disciformis on Acer, April 1979, H. J. M. Bowen (E, IMI 244538!). 

IX. LIBERTIELLA Speg. & Roum. 

in Spegazzini. Revue mycoi 2 : 22 (1880). 
Nicholsoniella Kuntze, Rev. Gen. PL 2 : 862 (1891). 
Shecutia Nieuwl., Am. Midi. Nat. 4 : 379 (1916). 

Conidiomata pyenidial, arising singly or in small groups, often aggregated, subglobose, 
superficial, whitish to pale orange, ostiole forming by a disintegration of the upper wall within 
an area delimited by a reddish-brown ring of tissue; walls composed of almost hyaline very 
thick-walled pseudoparenchymatous cells, often interspersed with orange granules. Conidio- 
phores hyaline, irregularly sympodially branched, short, cylindrical. Conidiogenous 
cells enteroblastic, phialidic, arising laterally or terminally from the conidiophores, rarely 
proliferating. Conidia adhering in gelatinous masses, ellipsoid to obovoid with a slightly 
truncated base, hyaline, simple, smooth-walled. 

Type species: Libertiella malmedyensis Speg. & Roum. 

Number of species: Monotypic. Other species previously referred to this genus are now 
placed in different genera (see Index). 

Observations: Both Nicholsoniella Kuntze and Shecutia Nieuwl. were introduced as new 
names for Libertiella in the belief that Libertiella was a homonym of the earlier Libertella 
Desm.; Nieuwland (1916) was evidently unaware of the new name introduced by Kuntze. 
However, as pointed out by Sutton (1977 : 113), in coining the new generic name Libertiella, 
Spegazzini and Roumeguere deliberately inserted the 7' and made it clear in a footnote that 
they were doing this to prevent confusion with Libertella Desm. The generic name 
Libertiella is consequently not to be treated as a later homonym of Libertella and can be 
retained. 

1. Libertiella malmedyensis Speg. & Roum., in Spegazzini, Revue mycol 2 : 22 (January 
1880). 
(Figs. 16A-B, 17A-E) 

Type: Belgium, "prope malmedyanum\ on Peltigera spuria (Ach.) DC, 'Hieme\ M.-A. Libert 
(BR — holotype, non vidi; K — isotype!). 
Nicholsoniella malmedyensis (Speg. & Roum.) Kuntze, Rev. Gen. PL 2 : 863 (1891). 
Shecutia malmedyensis (Speg. & Roum.) Nieuwl. Am. Midi. Nat. 4 : 379 (1916). 
Zythia peltigerae Lib. ex Cooke, Grevillea 8 : 83 (March 1 880), nom. illegit. (Art. 63). 
Type: Belgium, [Malmedy], on Peltigera spuria (Ach.) DC, M.-A. Libert (BR — holotype, non vidi; 
K — isotype!). 
Libertiella peltigerae (Lib. ex Cooke) Keissler, Rabenh. Krypt.-Fl. 8 : 582 (1930). 

Exsiccatae: Roumeguere, Fungi sel. Gall. no. 671 (K!). — Thiimen, Mycoth. univ. no. 1775 (K!). 

Conidiomata pyenidial, superficial, with only the base immersed in the host, arising 
singly or more commonly in groups, often aggregated, subglobose, whitish to pale orange, 
mainly (100-)150-200(-300)//m diam, becoming flattened at the apex with the develop- 
ment of the ostiole; ostiole forming after the pyenidial cavity is full of spores, demarcated by 
a limiting reddish-brown ring of tissue about 25 pm wide encircling the whitish 50-75 pm 
diam pore; pyenidial wall mainly 10-12 pm thick, composed of ± hyaline translucent 
thick-walled pseudosclerenchymatous cells, cells mainly 3-4*5 pm thick, walls of the inner 
layers of cells thinner and these cells often orientated perpendicularly to the outermost 
layer(s) which are often interspersed with minute orange granules also present on the wall 
surface. Conidiophores arising from the inner wall of the pyenidial cavity, cylindrical, 
irregularly sympodially branched, hyaline, smooth-walled, 2-8 pm tall and 3—3*5 pm 



LICHENICOLOUS COELOMYCETES 



31 







A^ 



i.^v^^^ 







v*-**- - " 



Fig. 16 Libertiella malmeydensis (K — isotype). A, Vertical section of pycnidium (x 500). 
B, Vertical section of pycnidial wall with conidiogenous cells and conidia (x 1020). 



wide. Conidiogenous cells enteroblastic, arising terminally and laterally from short conidio- 
phores or directly from the inner wall of the pycnidial cavity, cylindrical, phialidic with a 
short collarette, rarely proliferating, hyaline, 10-12 x 2-3*5 //m. Conidia abundant, 
adhering together in gelatinous masses, ellipsoid to obovoid with a slightly truncated base, 
rather irregular in outline, hyaline, simple, rarely guttulate, smooth-walled, (5— )6— 8 x (2-)3- 
4//m. 

Host: Peltigera spuria (Ach.) DC, thallus. The pycnidia arise most abundantly on the 
underside of the thallus but are also present in groups on the upper cortex, especially where 
soredia were formerly produced in the P. erumpens (Taylor) Vainio morph of this species. 
Libertiella malmedyensis causes relatively minor damage to infected thalli and does not 
lead to the formation of necrotic patches. The host of this fungus has consistently been cited 
as P. polydactyla (Necker) Hofifm., but Libert's material is P. spuria and not P. polydactyla. 

Distribution: Belgium and Poland. It was evidently very abundant in the type locality as in 
addition to sending material to several herbaria, Libert left enough in Brussels for the 
exsiccatae of both Roumeguere and de Thumen. 

Observations: Libertiella malmedyensis shows some similarity to Karsteniomyces 
peltigerae (p. 22) but differs in both the method of conidiogenesis and the non-septate and 
differently shaped ascospores; in addition the pycnidial wall in K. peltigerae has much more 
thickened pseudosclerenchymatous cells than L. malmedyensis and the conidia are not 
markedly gelatinous. 

Keissler (1930 : 582) took up the epithet ' peltigerae" for this species as he believed that was 
published in March 1880 while 'malmedyensis' came out in April 1880. The issue of the 
Revue mycologique including Spegazzini's paper was volume 2 part 1 as this has the title of 
his paper printed on the front wrapper together with 'Janvier 1880'. In Grevillea volume 8 
number 47, dated March 1 880 on the first page, and which included Cooke's article 'Relique 
Libertianae' on pp. 81-87, on p. 120 of this same number appears under k Cryptogamic 
Literature' the entry 'Roumeguere, C. Reliquae Libertianae, in "Revue Mycologique". Jan., 



32 



D. L. HAWKSWORTH 




Fig. 17 Libertiella malmeydensis (K — isotype). A, Pycnidia on the host thallus showing 
the deeply pigmented ring around the ostiole. B, Vertical section of pycnidium. C, 
Vertical section of pycnidial wall with granular encrustations on the outer cells. D, 
Conidiophores and conidiogenous cells. E, Conidia. 



1 880.' There can consequently be no doubt that Spegazzini's article, which was an appendix 
to that of Roumeguere, appeared before the printing and distribution of the part ofGrevillea 
including Cooke's paper. The epithet " malmedyensis" must consequently be reinstated for 
this species because it predates "peltigerae\ and in the absence of evidence to the contrary 
'malmedyensis' is taken to have been validly published in January 1880. Keissler's date for 
i malmedyensis i might have arisen from a hasty confusion of the wrappers of the Revue; 
volume 2 part 2 has 'Avril 1 880' printed on it. 

Roumeguere's exsiccatum included a description of Libertiella malmedyensis on the 
printed label. The fascicle including no. 671, fasc. VII, was also issued in January 1880 
according to a mention in Revue mycologique 2(1): 27. I have not been able to sequence the 



LICHENICOLOUS COELOMYCETES 33 

exsiccata and journal article within January 1880; it is even possible that they were 
distributed simultaneously to save postage as Roumeguere then edited the journal! 

Additional specimen: Poland: Puszcza Piska, on Peltigera spuria, June 1958, J. Zielinska (K!). 

X. LICHENOCONIUM Petrak & H. Sydow 

Beih. Repert. Spec. nov. Regni veg. 42 : 432 (1927). 

See Hawksworth (1977 : 170) and Sutton (1980 : 118) for modern descriptions of the 
genus. 

Type species: Lichenoconium lichenicola (P. Karsten) Petrak & H. Sydow. 

Number of species: Eight, all obligately lichenicolous. 

Observations: Two additional species, Lichenoconium boreale (P. Karsten) Petrak & H. 
Sydow and L. pertusariicola (Nyl.) D. Hawksw., were only retained in Lichenoconium by 
Hawksworth (loc. cit.) pending further research on the generic names proposed for 
Coelomycetes. From the major studies of Sutton (1977, 1980) it is evident that there is no 
currently acceptable genus available for either of these two fungi. L. pertusariicola is 
therefore referred to the new genus Laeviomyces above, and its separation from 
Lichenoconium is discussed under that name (p. 27). L. boreale is not a lichenicolous 
fungus but occurs on conifer lignum; that fungus differs from Lichenoconium in having a 
pycnidial wall which consists of interwoven hyphae (textura intricata), very slender 
conidiogenous cells, and almost colourless to pale olivaceous conidia which are ellipsoid to 
almost allantoid, clearly guttulate, with smooth walls when studied by light microscopy 
(although with a delicate close-packed verruculose ornamentation in SEM). The new generic 
name Xeroconium D. Hawksw.* is therefore introduced for this species here; a detailed 
description and illustrations of Xeroconium boreale (under the name Lichenoconium 
boreale) were presented in Hawksworth (1977 : 171-172, fig. 1A, pi. 22 A-E) and further 
drawings are given by Sutton (1980 : 1 19, fig. 56 A-C). 



Key to the species 

1 Pycnidia mainly exceeding 100 /im diam 

Pycnidia mainly less than 80 //mdiam 

2 Conidia mainly exceeding 4 jum in length, distinctly echinulate or verruculose 

- Conidia (2-)2-5-3'5(-4)x2-3 //m, rather smooth-walled; pycnidia (60-)80-120(-150)x 

50-80(-100)//m Lichenoconium pyxidatae (p. 36) 

3 Conidia globose to subglobose, sometimes rather angular 

Conidia ellipsoid with a tapering and abruptly truncated base, (4-)6-8(-9)x 3^1(-6)//m; 

pycnidia 100-200 fim diam; conidiogenous cells (6— )8— 1 3(— 1 5) x 2-3'5(^*5) //m 

Lichenoconium lichenicola (p. 36) 



4 Conidia (2-5-)3-4'5(-6)/im diam; pycnidia (80-)100-175(-200)//m diam; conidiogenous 

cells (5-)6-8(-l l)x(2-)2-5^//m Lichenoconium xanthome (p. 37) 

Conidia 5-7(-7'5)x 3'5-5(-6)//m; pycnidia 100-175(-200)/im diam; conidiogenous cells 
(7— )8— 10(— 12) x (2*5— )3— 3-5 fim Lichenoconium cargillianum (p. 35) 

5 Pycnidia mainly exceeding 50 jum diam; conidia mainly exceeding 3*5 /um diam . 

- Pycnidia (20-)30-50(-60) fim diam; conidiogenous cells (35-)4-5(-6)x (2-)3-35(-4)//m; 

conidia 2-3 , 5H) J um Lichenoconium erodens (p. 35) 

*Xeroconium D. Hawksw. gen. nov. 

Genus ad Coelomycetes Sphaeropsidales pertinens. Similis generi Lichenoconio Petrak & H. Sydow sed diftert in 
muris pycnidiis textura intricata, cellulis conidiogenis gracilioribus, et conidiis ellipsoideis ad plusminusve 
allantoideis, subhyalinis ad pallide olivaceis, gutuilatis, laevibus (sed per microscopium electronicum minuto- 
verruculosis). — Species holotypica, adhuc unica, est Xeroconium boreale (P. Karsten) D. Hawksw. comb. nov. 
(basionym: Levieuxia borealis P. Karsten, Hedwigia 26 : 126, 1887). 



34 



D. L. HAWKSWORTH 

B 




lOjLim 



Fig. 18 Lichenoconium species, conidiogenous cells and conidia. A, L. echinosporum 
(UPS — holotype). B, L. erodens (herb. Christiansen — holotype). C, L. cargillianum 
(E— holotype). D, L. lichenicola (H-KARST 1246— holotype). E, L. lecanorae (IMI 192264). F, 
I. pyxidatae (L — holotype). G, L. usneae (K — isotype). H, L. xanthoriae (C — holotype). 
Reproduced from Hawksworth ( 1977 : 162, 164). 



LICHENICOLOUS COELOMYCETES 35 

6 Conidiogenous cells mainly exceeding 7 jum tall 7 

- Conidiogenous cells (4-)5-7(-8)x(2-)3-3'5(-4)//m; conidia (2-5-)3-4'5(-5'5) i um; pyc- 
nidia (30-)40-80(- 100) //m diam Lichenoconium lecanorae (p. 36) 

7 Conidia verruculose, (2'5-)3^1(-5)//m diam; pycnidia (40-)50-80(-100)//m diam; 

conidiogenous cells (5— )7— 9(— 1 1 ) x (2-)2- 5-3 -5(^1) pim . . Lichenoconium usneae (p. 37) 
Conidia sparsely echinulate, 4*5-5 x 3-4 //m; pycnidia 65-125 /zm diam; conidiogenous 
cells (7— )8— 10(— 1 l)x 2-3 //m Lichenoconium echinosporum (p. 35) 

1. Lichenoconium cargillianum (Lindsay) D. Hawksw., Persoonia 9 : 172 (1977). 
(Fig. 18C) 

See Hawksworth (1977 : 172-3) for description, illustrations and synonymy. 

Hosts: Parmelia perforata Ach., Ramalina yemensis (Ach.) Nyl. and Usnea florida (L.) 
Wigg., predominantly in apothecia. 

Distribution: British Isles, Mexico and New Zealand. 

2. Lichenoconium echinosporum D. Hawksw., Persoonia 9 : 1 73 (1977). 
(Fig. 18 A) 

See Hawksworth (1977 : 173-174) for description and illustrations. 

Host: Heterodea muelleri (Hampe) Nyl., a pathogen on the thallus. 

Distribution: Australia. I have only seen the holotype but Filson (1978 : 17) reports seeing 
pale brown necrotic patches like those due to this fungus from a wide range of Australian 
localities. 

3. Lichenoconium erodens M. S. Christ. & D. Hawksw., in Hawksworth, Persoonia 9 : 174 

(1977). 
(Fig. 18B) 

See Hawksworth (1977 : 174-177) for description and illustrations. 

Hosts: Cladonia coniocraea (Florke) Sprengel, C. digitata (L.) HofTm., Evernia prunastri 
(L.) Ach., Hypogymnia bitteriana (Zahlbr.) Krog, H. physodes (L.) Nyl., Lecanora chlarona 
(Ach.) Nyl., L. conizaeoides Nyl. ex Crombie, Parmelia afrorevoluta Krog & Swinscow (syn. 
P. revoluta auct. p.p.), P. caperata (L.) Ach., P. crinita Ach., P. galbina Ach., P. laevigata 
(Sm.) Ach., P. perlata (Huds.) Ach., P. saxatilis (L.) Ach., Parmeliopsis ambigua (Wulfen) 
Nyl., Pertusaria hymenea (Ach.) Schaerer and P. pertusa (Weigel) Tuck. 

Distribution: Austria, British Isles, Denmark, France, Germany, Sweden and the U.S.A. 

Observations: A wide-ranging species which has been found on seven additional hosts 
since its original description (Hawksworth, 1977). By far the most pathogenic species of the 
genus, typically producing bleached areas on thalli which subsequently die. In cases where 
this and other species of the genus are able to occur on the same host species, the symptoms 
generally differ, Lichenoconium erodens always causing the most damage. This is also true 
for the additional hosts included in the list above, including the Cladonia species and 
Lecanora conizaeoides: its effects on the latter are illustrated and compared with those due 
to L. lecanorae by Christiansen ( 1 980). 

Specimens (additional to those listed in Christiansen, 1980; Hawksworth, 1977, 1980c; and 
Hawksworth & Minter, 1980): Austria: Steiermark, Grazer Bergland, Graz, Maria Trost, on 
Hypogymnia physodes, 24 March 1974, J. Hafellner 568 (GZU!); Steiermark, Frauenalm S. of Musau, 
on Cladonia digitata, 18 June 1978, J. Hafellner 3897 (GZU!). —British Isles: England, Middlesex, 
Ruislip, Park Wood, on Cladonia coniocraea on Quercus, 9 December 1979, D. L. Hawksworth 5009 
(IMI 244544a!); S. Devon, Cornwood, Dendles Wood, on Parmelia laevigata, 29 February 1976, D. L. 
Hawksworth 4293b (IMI 201661/?!); S. Devon, Okehampton, nr Fatherford Viaduct, on Hypogymnia 



36 D. L. HAWKSWORTH 

physodes on Quercus, 30 March 1979, D. L Hawksworth 4903 (IMI 236902!); Dorset, Studland Heath, 
on // physodes on Betula. 25 June 1979, D. L Hawksworth 4952 (IMI 240076!); Hampshire, New 
Forest, Stubbs Wood, on H. physodes, 13 May 1979, D. L. Hawksworth 4936 (IMI 238699!); S. 
Somerset, nr Stoke Perol, on H. physodes on Quercus, 6 October 1979, D. L Hawksworth 5002 (IMI 
243338!); S. Somerset, Porlock, Horner Combe, on P. laevigata on Betula, 21 September 1980, D. L. 
Hawksworth 5073 (IMI 251491!); Scotland, Dunbartonshire, Loch Lomond, Ross Park, on P. caperata 
on Quercus. 9 October 1978, B. J. Coppins 3688 (E, IMI 232973!); Wales, Anglesey, Abberfraw sand 
dunes, on Evernia prunastri, 1 7 October 1977, D. L Hawksworth 465 1 (IMI 2 1 7423!). 

4. Lichenoconium lecanorae (Jaap) D. Hawksw., Bull. Br. Mus. nat. Hist. (Bot.) 6: 183 

(1979). 
(Fig. 18E) 

See Hawksworth (1977 : 178-182, sub L. parasiticum D. Hawksw.) for description and 
illustrations. 

Hosts: Evernia prunastri, Lecanora admontensis Zahlbr., L. carpinea (L.) Vainio, L. 
chlarotera Nyl., L. conizaeoides Nyl. ex Crombie, L. pallida (Schreber) Rabenh., L. 
polytropa (HofTm.) Rabenh., L. subfuscata Magnusson, L. superfluens Magnusson, Parmelia 
barren (Sm.) Turner, P. galbina Ach., P. pastillifera (Harm.) R. Schub. & Klem., P. saxatilis 
(L.) Ach., P. sulcata Taylor, Rhizoplaca chrysoleuca (Sm.) Zopf and Squamarina lentigera 
(Weber) Poelt. There are also some unconfirmed literature reports (see Hawksworth 
1977:179). 

Distribution: Austria, ?Belgium, British Isles, Canada, Czechoslovakia, Denmark, France, 
Germany, Hungary, Italy, Netherlands, Spain, Sweden, Switzerland and the U.S.A. 

Observations: The symptoms of this species on Lecanora conizaeoides are illustrated and 
compared with those due to Lichenoconium erodens by Christiansen (1980). 

Specimens (additional to those listed by Hawksworth, 1977): Austria: Steiermark, Schladminger 
Tauren. Kleinsolk-Obertal, on Lecanora polytropa, 12 August 1974, J. Hafellner 3932 (GZU!). 
— British Isles: England, Hampshire, New Forest, Stubbs Wood, on Parmelia sulcata, 13 May 1979, D. 
L Hawksworth 4937 (IMI 238700!); S. Devon, Slapton Ley Nature Reserve, Peasdish, on P. sulcata, 23 
August 1980, D. L Hawksworth 5044b (IMI 251 1262!); loc. cit., on Lecanora sp., 23 August 1980, D. L 
Hawksworth 5044a (IMI 251261!). — France: Dept. Isere, Rhone-Tal E. Lyon, on Squamarina 
lentigera, 13 July 1975, J. Hafellner 1575 (GZU!). —Italy: Sudtirol, Mendelgebirge, Penegal SW. von 
Bozen. on Lecanora carpinea, 20 October 1975, J. Hafellner 1092 (GZU!); Otztaler Alpen, Vinschgau, 
on Rhizoplaca chrysoleuca, 18 October 1975, J. Hafellner 991 p.p. (GZU!). 

5. Lichenoconium lichenicola (P. Karsten) Petrak & H. Sydow, Beih. Repert. nov. Spec. Regni 

veg. 42: 432(1927). 
(Fig. 18D) 

See Hawksworth (1977 : 177-178) for description and illustrations. Further drawings are 
provided by Sutton (1980 : 1 19 fig. 55a-c). 

Host: Physcia aipolia (Ehrh. ex Humb.) Fiirnrohr, apothecia. 

Distribution: Finland. Known only from the type collection. 

6. Lichenoconium pyxidatae (Oudem.) Petrak & H. Sydow, Beih. Repert. nov. Spec. Regni 

veg. 42:435(1927). 
(Fig. 18F) 

See Hawksworth (1977 : 184-1 85) for description and illustrations. 

Hosts: Cladonia arbuscula (Wallr.) Rabenh., C. cenotea (Ach.) Schaerer, C.fimbriata (L.) 
Fr., C. incrassata Florke, C. pocillum (Ach.) O.-J. Rich., and C. pyxidata (L.) Hoffm.; 
usually on the podetia but occasionally on squamules. There are also unconfirmed reports 
from additional hosts in Cladonia and other genera, the latter certainly erroneous (see 
Hawksworth, loc. cit.). 



LICHENICOLOUS COELOMYCETES 37 

Distribution: Austria, British Isles, Denmark, France, Germany, the Netherlands and 
Sweden. 

Observations: Harmand (1907 : 226) stated that Cladonia rangiferina f. moribunda Harm, 
was based on deformed podetia with verrucae caused by a fungus close to Lichenoconium 
pyxidatae but 4 il se distingue par des spores franchement brunes et ovoides' according to 
Vouaux who Harmand indicates studied the material. I have not seen type or authentic 
material ofthistaxon. 

7. Lichenoconium usneae (Anzi) D. Hawksw., Persoonia 9 : 185 (1977). 
(Fig. 18G) 

See Hawksworth (1977: 185-1 90) for description, illustrations and synonymy. 

Hosts: Anaptychia ciliaris (L.) Korber, Bryoria fuscescens (Gyelnik) Brodo & D. Hawksw., 
Cladonia arbuscula (Wallr.) Rabenh., C. cariosa (Ach.) Sprengel, Hypogymnia physodes (L.) 
Nyl., Lecanora pacifica Tuck., Letharia vulpina (L.) Hue, Parmelia conspersa (Ehrh. ex 
Ach.) Sprengel, P. exasperata (Ach.) de Not., P. glabratula (Lamy) Nyl., P. olivacea (L.) 
Ach., P. pulla Ach., P. rudecta Ach., P. saxatilis (L.) Ach., P. verruculifera Nyl., Physcia 
aipolia (Ehrh. ex Humb.) Fiirnrohr, P. stellaris (L.) Nyl., Physconia pulverulacea Moberg, 
Ramalina cf. baltica Lettau, R. calicaris (L.) Fr., R.fraxinea (L.) Ach., R. siliquosa (Huds.) 
A. L. Sm., R. subgeniculata Nyl., Usneafilipendula Stirton aggr., and U. cf.florida (L.) Wigg. 
For unconfirmed literature reports see Hawksworth (loc. cit.). 

Distribution: Austria, British Isles, Canada, Canary Islands, Czechoslovakia, Denmark, 
France, Germany, Italy, Norway, Spain, Sweden, Switzerland, Yugoslavia (Keissler, 
1933: 388) and the U.S.A. 

Specimens (additional to those listed in Hawksworth, 1977): Austria: Steiermark, Stubalpe, Gaberl, 
on Usnea cLjlorida, 19 December 1976, J. Poelt & J. Hafellner 1825 (GZU!); Steiermark, Kreuzberg, 
on Cladonia sp., 16 March 1975, W. Moschl & H. Pittoni [herb. Hafellner 437] (GZU!); Nockgruppe, 
Afritzer Berge, Karten, on Usnea sp., 27 August 1974, J. Poelt (GZU!). — British Isles: England, S. 
Devon, Slapton, Duck Marsh, on Ramalina fraxinea, 5 September 1978, D. L. Hawksworth 4869 (IMI 
231752!); Scotland, Midlothian, Carlops, Hobbies House, on Bryoria fuscescens, 29 March 1977, B. J. 
Coppins 2727 (E!). — Canada: Alberta, Kananaskis Range, between Wind Ridge and Mount Longhead, 
on Letharia vulpina, 22 June 1978, D. C Lindsay (IMI 229869/?!). —France: Dept. Vas, He de Port 
Cros, on Ramalina calicaris, 17 July 1973, Y. Rondon [herb. Hafellner 1826] (GZU!). 

8. Lichenoconium xanthoriae M. S. Christ., FriesiaS : 212 (1956). 
(Fig. 18H) 

See Hawksworth (1977: 1 90-1 92) for description and illustrations. 

Hosts: Cetraria sepincola (Ehrh.) Ach., Cetrelia olivetorum (Nyl.) Culb. & C. Culb. s. lat., 
Xanthoria parietina (L.) Th. Fr., X. polycarpa (Hoffm.) Rieber. The collections on Cetrelia 
have been reported on by Hawksworth & Minter ( 1 980). 

Distribution: British Isles, Czechoslovakia, Denmark and Sweden. 
XI. LICHENODIPLIS Dyko & D. Hawksw. 
in Hawksworth & Dyko, Lichenologist 11 : 5 1 (1979). 

See Hawksworth & Dyko (1979) for further information on this genus. 
Type species: Lichenodiplis lecanorae (Vouaux) Dyko & D. Hawksw. 
Number of species: Two, both obligately lichenicolous. 

Key to the species 

1 Conidiomata 50-120 //m diam; conidia 4-75 x 2-3 fim . Lichenodiplis lecanorae (p. 38) 

- Conidiomata 50-70 /zm diam; conidia 95—1 3 x 4-4-5 ^m . . Lichenodiplis lichenicola (p. 38) 



38 D. L. HAWKSWORTH 

1. Lichenodiplis lecanorae (Vouaux) Dyko & D. Hawksw., in Hawksworth & Dyko, 

Lichenologist 11: 52(1979). 

See Hawksworth & Dyko ( 1 979) for description, illustrations and synonymy. 

Hosts: Caloplaca caesiorufa (Wibel) Flagey, C. cerina (Ehrh. ex Hedw.) Th. Fr., C. 
ferruginea (Huds.) Th. Fr., C. flavovirescens (Wulfen) Dalla* Torre & Sarnth., Evernia 
prunastri (L.) Ach., Lecanora confusa Almb., L. dispersa (Pers.) Sommerf., L. pallida 
(Schreber) Rabenh., L. polvtropa (Hoffm.) Rabenh., L. saligna (Schrader) Zahlbr., Pertusaria 
albescens (Huds.) M. Choisy & Werner and P. cf. leioplaca DC. Hawksworth & Dyko (1979) 
also note that there are literature records from Caloplaca lactea (Massal.) Zahlbr., Lecanora 
expallens Ach., L. varia (Hoffm.) Ach., Diploschistes scruposus (Schreber) Norman, 
Lecidella elaeochroma (Ach.) M. Choisy, Lecidea enteroleuca Ach., Micarea nitschkeana 
(Lahm ex Rabenh.) Harm, and unidentified Lecidea and Pertusaria species. 

Distribution: Hawksworth & Dyko (1979) saw material from the British Isles, Canary 
Islands, Denmark, France, Germany, Spain and Sweden; they also noted a literature report 
from the U.S.S.R. (Novaya Zemyla). Also mentioned from Morocco by Werner (1972 : 95) 
on Pertusaria pustulata f. glabrata (Anzi) Hue. 

Observations: While examining material in H-NYL in August 1979, I discovered that 
Nylander had given this species the herbarium name "Dichaena pertusariae'; this name does 
not, however, ever appear to have been used in print. 

Specimens (additional to those listed by Hawksworth & Dyko, 1979): British Isles: England, S. 
Devon, Torquay, on Pertusaria cf. leioplaca, [?R. Deakin] [Leighton, Lich. exs. no. 245; sub 
Opegrapha atra var. parella.] (E!); Westmorland, Brathay church, on Lecanora cf. saligna, 22 
September 1979, A. Henderson (herb. Henderson!). Scotland, W. Inverness, Strontian, Horsley Hall, on 
Pertusaria albescens on Ulmus, 14 July 1966, P. W. James (IMI 237277!). — France: Brittany, Brest, on 
Pertusaria cf. leioplaca, Crouan 135 (H-NYL p.m. 7683!). 

2. Lichenodiplis lichenicola Dyko & D. Hawksw., in Hawksworth & Dyko, Lichenologist 

11 : 56(1979). 

See Hawksworth & Dyko (1979) for description, illustrations and a possible synonym. 

Host: Rinodina septentrionalis Malme, apothecia. 

Distribution: Hawksworth & Dyko (1979) indicated that the type and only known 
collection of this species came from Finnmark. However, Dr P. M. Jorgensen (in litt.) 
has kindly pointed out that the type almost certainly came from the Yenisey region of the 
U.S.S.R. 

Observations: I have also seen a further collection that may belong to this species (France: 
Vosges, Docelles, on Pertusaria cf. leioplaca, J. Harmand, herb. Vouaux!), with conidia 
9-12 x 4-5 pm, but hesitate to refer it definitely to Lichenodiplis lichenicola as the conidia 
are rather thick-walled, much darker brown, and tend to be somewhat broader in relation to 
their length. Until more collections are available which enable the variability of L. 
lichenicola to be determined, it would be premature to either describe this material as new or 
state that it definitely belonged here. 

XII LICHENOSTICTA Zopf 

Nova Ada Acad. Caesar. Leap. Carol. 70(4) : 263 (1898). 

Conidiomala pycnidial, arising singly, uniloculate, subglobose to broadly pyriform, 
erumpent to largely superficial, translucent brown to dark brown or black; walls composed 
of a few layers of intertwined brownish irregular hyphae forming a textura intricata, lined by 
a hyaline almost pseudoparenchymatous layer of cells, thickened near the ostiole. 
Conidiophores hyaline, irregularly branched, flexuous, multi-septate, arising from the inner 



LICHENICOLOUS COELOMYCETES 39 

cells of the pycnidial wall. Conidiogenous cells enteroblastic, phialidic, sometimes 
proliferating, integrated into chains, acro-pleurogenous, conidia arising from the apex of a 
chain and (for other conidiogenous cells) laterally, sometimes polyphialidic, hyaline, 
cylindrical to doliiform, channel and collarette distinct, narrow proliferating tissue often 
evident. Conidia adhering in slime and extruded as a gelatinous drop, lacriform, distinctly 
attenuated at the base, hyaline, simple, smooth-walled. 

Type species: Lichenosticta alcicornaria (Lindsay) D. Hawksw. (syn. L. podetiicola Zopf). 

Number of species: Monotypic. 

Observations: The generic name Dendrophoma Sacc. cannot be used for this species as it is 
a synonym of Dinemasporium Lev., a very different fungus (Sutton, 1968). The catenate 
arrangement of the conidiogenous cells and details of conidiogenesis show a strong similarity 
to those of the pycnidia of some lichenized members of the Lecanorales (see Vobis & 
Hawksworth, 1981). The normal pycnidia of Cladonia species have much more regularly 
branched conidiophores and mainly ellipsoid to bacillariform conidia. 

1. Lichenosticta alcicornaria (Lindsay) D. Hawksw., in Hawksworth et al, Lichenologist 
12: 107(1980). 
(Fig. 19A-G) 

Microthelia alcicornaria Lindsay, Q. Jlmicrosc. Sci. II, 9 : 349 (1869). 

Type: British Isles, England, Shropshire, Haughmond Hill, on Cladonia foliacea (Huds.) Willd., 
W. A. Leighton [Lich. exs. no. 15] (E — lectotype!; BM, E — isolectotypes!). 
Dendrophoma alcicornaria (Lindsay) Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 283 (1914). 
Lichenosticta podetiicola Zopf, Nova Acta Acad. Caesar. Leop. Carol. 70(4) : 263 (1898). 
Dendrophoma podetiicola (Zopf) Keissler, Ost. bot. Z. 60 : 62 (1910). 
Aposphaeria cladoniae Allescher & Schnabl, in Allescher, Ber. Bayer, bot. Ges. 4 : 32 (1 896). 

Type: Germany, Munich, Pullach, on Cladonia fimbriata (L.) Fr., October 1894, J. N. Schnabl 
(M — holotype!). 
Phoma cladoniae (Allescher & Schnabl) Keissler, Annln naturh. Mus. Wien 34 : 77 ( 1 92 1 ). 
? Aposphaeria cladoniae var. floerkeanae Vouaux, in Harmand, Lich. Fr. 3 : 335 (1907). 

Exsiccatae: Leighton, Lich. exs. no. 1 5 (sub Cladonia alcicornis; BM!, E!). — Rasanen, Lich. fenn. exs. 
no. 600 (sub Dendrophoma alcicornaria*; BM!). 

Icones: Lindsay, Trans. R. Soc. Edinb. 22: pi. 8 fig. 3 (1859). — Zopf, Nova Acta Acad. Caesar. Leop. 
Carol. 70(4) : 264 figs 22-23, 265 figs 24-25 (1898). 

Conidiomata pycnidial, immersed or sometimes becoming to ] erumpent, arising singly, 
often numerous, scattered, translucent brown to dark brown or black, sometimes shiny, 
(50-)80-120(-150)//m, subglobose to broadly pyriform, ostiolate; pycnidial wall irregular 
in outline, mainly 5-10 //m thick but to 20-25 pm thick around the ostiole, composed of 
1-3 (or more near the ostiole) irregular layers of intertwined hyphae, hyphae olivaceous 
brown to dark brown, uneven in diameter, very variable in shape, sometimes strongly in- 
flated between the septa, 3-7 jum wide, thick-walled near the ostiole, inner cells subhyaline 
and almost pseudoparenchymatous in parts. Conidiophores hyaline, irregularly branched, 
occupying the whole of the pycnidial cavity in young pycnidia, flexuous, multi-septate, 
arising from the inner cells of the pycnidial wall. Conidiogenous cells enteroblastic, 
phialidic, sometimes percurrently proliferating, integrated into chains, acro-pleurogenous 
with conidia arising from the apex of a chain and (for other conidiogenous cells) laterally or 
near a transverse septum, sometimes polyphialidic, hyaline, cylindrical to doliiform, mainly 
2*5^ pm wide and 4-6 jum long, channel and collarette distinct, this (and often 

*No. 350, distributed by Rasanen under this name, is not this species but probably an unknown pycnothyriaceous 
fungus. Mentioned by Sandstede (1931) as present in his Clad. Exs. nos. 601, 870-2, and 1574 but these numbers 
have not been studied by me. 



40 



D. L. HAWKSWORTH 








I 



lmm 



50/im 






Fig. 19 Lichenosticta alcicornaria. A-E (Rasanen, Lick. Exs. no. 600, K); A, pycnidia on 
podetia of the host; B, vertical section of pycnidium; C, cells from the pycnidial wall; D, 
chains of conidiogenous cells; E, conidia. F (Hafellner 1834), Conidiogenous cells and 
conidia. G (BM — isotype), Conidiogenous cells and conidia. 



conspicuous narrow proliferations) 1-1 "5 /mi wide. Conidia adhering in slime and extruded 
as a gelatinous drop, lacriform, plano-convex to concavo-convex, rounded at the apex and 
distinctly attenuated at the base, hyaline, simple, often 1 or more guttulate, smooth-walled, 
(6-)6-5-l 0(-l 1 ) x (2-)3-4-5(-6) //m. 



LICHENICOLOUS COELOMYCETES 41 

Hosts: I have examined material on Cladonia arbuscula (Wallr.) Rabenh. (C sylvatica 
auct.), C fimbriata (L.) Fr., C foliacea (Huds.) Willd. and C gracilis (L.) Willd. Also 
mentioned from C cornuta (L.) Hoffm. by Keissler (1910:2), C pyxidata (L.) Hoffm. by 
Keissler (1930 : 551); from C acuminata (Ach.) Norrlin (C norrlinii Vainio), C. portentosa 
(Dufour) Coem. (C impexa Harm.), and C ciliata var. tenuis (Florke) Ahti by Sandstede 
(1931 : 52, 76, 340); and from C coniocraea auct. by Folan & Mitchell (1970 : 169). The 
fungus occurs on both podetia and squamules (especially the lower surfaces) but causes little 
damage so is easily overlooked. Probably parasymbiotic. 

Distribution: Austria (Keissler, 1930 : 551), the British Isles (England; Ireland, Folan & 
Mitchell, 1970 : 169), Canada (Newfoundland; Keissler, 1910:2), Finland, Germany, Italy 
and Sweden (Keissler, 1910:2; Santesson, 1949 : 142). 

Observations: The fungus is easily distinguished from other Coelomycetes on Cladonia by 
the characteristic lacriform conidia and catenately arranged acro-pleurogenous conidio- 
genous cells. Further, it never appears to induce the formation of gall-like growths. 

The conidial shape is rather constant in this species but there is a considerable degree of 
variation in the sizes of conidia between different collections. For example, on a specimen of 
C. arbuscula (IMI 232629!) they measured (8— )9— 10(— 1 1) x 4*5-6 //m while on one of C 
gracilis {Lick Fenn. Exs. no. 600, BM!) they were (6— )6'5— 8(— 8 5) x 3^(-5)/zm (see Fig. 
19E-G). It is thus conceivable that more than a single species might be involved, but in the 
current state of our knowledge of the variability of Lichenosticta alcicornaria it would be 
imprudent to attempt to recognise two taxa at the present time. The types of both 
Aposphaeria cladoniae and Microthelia alcicornaria have conidia in the lower size range, as 
did that of Lichenosticta podetiicola according to the measurements given by Zopf 
(1898:264). 

Lindsay (1869& : 349) did not provide a description of this species but the name is validly 
published because he refers to his earlier work (Lindsay 1859 : 161), where a description and 
illustrations were provided; his epithet must therefore be adopted as it pre-dates that of Zopf. 
No authentic material of Lichenosticta podetiicola could be traced in B (B. Hein, in litt.) or 
M (H. Hertel, in litt.), but the figures published by Zopf (1898 : 264-5) leave little doubt the 
taxa are conspecific, even though Zopf did not show any conidiogenous cells, as the conidia 
are also of the correct shape and size. 

Aposphaeria cladoniae var.floerkeanae could not be located amongst Harmand's material 
in Angers (M. Guerlesquin, in litt.) and is not represented in the remnants of Vouaux's 
herbarium (Rondon, 1970). It was distinguished by conidia 8 x 3 jum so is tentatively 
retained here as those dimensions are ± within the range of Lichenosticta alcicornaria as 
interpreted here. 

Additional specimen: Italy: Siidtirol, Siidtiroler Dolomiten, Passo di Rolle, S Abhange des Mt 
Castellazzo, on Cladonia arbuscula, 23 October 1976, J. Hafellner 1834 (herb. Hafellner!, IMI 
232629!). 



XIII. MICROCALICIUM Vainio [ANAMORPHS] 

Acta Soc. Fauna Flora fenn. 57(1) : 77 (1927). 

Conidiomata pycnidial, arising singly or in small groups on the thallus of the host, 
subglobose, erumpent to almost sessile, with the base to half immersed, dark brown to almost 
black, ostiolate; walls composed of several layers of pseudoparenchymatous cells, cells in the 
upper parts of the pycnidium very thick-walled, olivaceous, aeruginose or brown, almost 
sclerenchymatous, cells in the lower parts of the pycnidium similar or sometimes subhyaline 
to pale brown, thin-walled, irregular in shape and often almost hyphal in parts. 
Conidiophores absent. Conidiogenous cells enteroblastic, lining the pycnidial cavity, 
acrogenous, subcylindrical to cylindrical, phialidic, with a particularly distinctive often 
elongate canal, not proliferating, hyaline, smooth-walled. Conidia arising singly, not 



42 D. L. HAWKSWORTH 

catenate, subglobose to narrowly ellipsoid, sometimes slightly apiculate at the base, hyaline, 
simple, thin-walled, smooth-walled, extruded through the ostiole in a mucilaginous drop. 

Type species: Microcalicium subpedicellatum (Schaerer) Tibell (syn. M. disseminatum 
(Ach. ex Fr.) Vainio). 

X umber of species: Four species of Microcalicium (Caliciales) were accepted by Tibell 
(1978) in a critical revision of the genus. Anamorphs are known for two of the species and 
only these are treated here. Tibell's work should be consulted for information on the 
teleomorphs. 

Observations: No suitable coelomycete generic name could be found for the anamorphs of 
Microcalicium species. I have not introduced a separate generic name for them here because 
they often (though not exclusively) occur together with the teleomorph, and, when found 
separately, 'anamorph' can easily be added after the name. Further, as the genus includes 
lichenized and not only lichenicolous fungi, Art. 59.1 should perhaps be invoked to prevent 
the introduction of an anamorphic name, as this rule, which permits a dual nomenclature in 
fungi with pleomorphic life-cycles, specifically excludes (correctly in my view; cf. 
Hawksworth, 1978/?: 236-237) lichen-forming taxa. 

ke\ to the species 

1 Conidia subglobose, 2-3 x 1-5-2 jum\ conidiogenous cells 6-9 x 2-3 pm 

Microcalicium subpedicellatum [anamorph] (p. 42) 
Conidia narrowly ellipsoid, 2-3 (-4) x 1*5-2 pm\ conidiogenous cells 3-5(-6)x 1 -5-2-5 
(-3)//m Microcalicium conversum [anamorph] (p. 42) 

1 . Microcalicium conversum Tibell, Bot. Notiser 131 : 237 (1978) [anamorph]. 
(Fig.20G-H) 

fcones: Tibell, Bot. Notiser 131 : 242 fig. 7B (1978) [anamorph]. 

Conidiomata pycnidial, arising singly, subglobose, erumpent to almost sessile or partly 
immersed, dark brown to black, 40-60(-80) pm wide and 50-80 //m tall; walls 8-12 jum 
thick, composed of several layers of reddish brown very thick-walled almost scleren- 
chymatous pseudoparenchymatous cells, becoming somewhat olivaceous in postassium 
hydroxide, 3-5 pm diam, basal wall apparently similar to the exposed parts at least in 
almost superficial pycnidia. Conidiophores absent. Conidiogenous cells enteroblastic, lining 
the pycnidial cavity, acrogenous, subcylindrical, phialidic, with a distinct and often elongate 
canal, not proliferating, hyaline, smooth-walled, 3-5 (-6 ) x l-5-2'5(-3)//m. Conidia arising 
singly, not catenate, narrowly ellipsoid, rounded at the ends, hyaline, simple, thin-walled, 
smooth-walled, sometimes guttulate, 2-2>(-A) x ( 1 — ) 1 • 5—2 jum, extruded through the ostiole 
in a mucilaginous drop. 

Hosts: Calicium viride Pers. and Chaenotheca subroscida (Eitner) Zahlbr., thalli. 

Distribution: The species was recorded by Tibell (loc. cit.) only from Argentina (Tierra del 
Fuego). Australia (Tasmania) and Chile (Magallanes). 

Observations: The anamorph of Microcalicium conversum is clearly separated from that of 
M. subpedicellatum on the basis of the more complete pycnidial wall, its reddish brown 
colour, the length of the conidiogenous cells, and the quite different shape of the conidia. 
The teleomorph also differs from that of M. subpedicellatum in having a reddish brown (not 
greenish) excipulum, and 1 (not 3-7)-septate ascospores (fide Tibell, loc. cit.). 

Specimen: Chile: Terr. Magallanes: Rio Rubens, near Hotel Rio Rubens (about 50 km SE of 
Natales), on ( 'alicium viride on Nothofagus pumila, 1 3 January 1 94 1 , R. Santesson 56 1 8 p.p. (S!). 

2. Microcalicium subpedicellatum (Schaerer) Tibell, Bot. Notiser 131:240 (1978) 

[anamorph]. 
(Fig. 20A-F) 



LICHENICOLOUS COELOMYCETES 



43 



1QQOft(L«°P.0Q O 




OqQ 

%0 




10/im 






Fig. 20A-F, Microcalicium subpediccllatum [anamorph] {Hafellner 1095): A, pycnidia on 
the host thallus; B, vertical section of pycnidium; C, cells from the upper part of the 
pycnidial wall; D, cells from the lower part of the pycnidial wall; E, conidiogenous cells: 
F, conidia. G-H, M. conversum [anamorph] (Santcsson 561H p.p); G, conidiogenous cells; H, 
conidia. 



44 D. L. HAWKSWORTH 

Icones: Tibell, Bot Notiser 131 : 245 fig. 10B, 246 fig. 1 1 (1978) [anamorph]. 

Conidiomata pycnidial, subglobose, arising singly or in small groups, erumpent with the 
lower \-\ remaining immersed, black, 40-60 pm diam; walls in the upper parts mainly 
10-1 5 um thick, composed of several layers of somewhat aeruginose very thick-walled 
almost sclerenchymatous pseudoparenchymatous cells, becoming brownish in potassium 
hydroxide, 3-5 urn diam, walls in the lower parts 5-8 jum thick, composed of 2-3 layers of 
thinner-walled subhyaline cells forming an irregularly pseudoparenchymatous to in places 
almost hyphal tissue. Conidiophores absent. Conidiogenous cells enteroblastic, lining the 
pycnidial cavity, acrogenous, subcylindrical to cylindrical, phialidic, with a distinctly 
thickened and elongate canal, not proliferating, hyaline, smooth-walled, 6-9 x 2-3 jum. 
Conidia arising singly, not catenate, subglobose, sometimes slightly pointed at the base, 
hyaline, simple, thin-walled, smooth-walled, 2-3 x l'5-2//m, extruded through the ostiole 
in a milky-white mucilaginous drop. 

Hosts: Calicium glaucellum Ach., C. viride Pers., Chaenotheca brunneola (Ach.) Mull. 
Arg., C. chrysocephala (Turner ex Ach.) Th. Fr., C. subroscida (Eitner) Zahlbr., C trichialis 
(Ach.) Th. Fr., and undetermined Caliciales, thalli. A parasitic species in which the pycnidia 
colonize at an early stage with to 15 per host areole. Infected thalli are eventually 
decolourized and may turn brownish with the formation of extensive lesions in the colonies. 
Sometimes apparently growing directly on lignum or with other algae, perhaps persisting in 
situ after the loss of the host in such cases. 

Distribution: Circumboreal. Recorded by Tibell (loc. cit.) from Austria, the British Isles 
(Scotland), Canada, France, Italy, Norway, Sweden, Switzerland and the USSR. 

Observations: The separation of this species from the anamorph of M. conversum is 
discussed under that species above. 

Specimens: Austria: Steiermark, Ostalpen, Gurktaler Alpen, Frauenalm S von Murau, on 
Chaenotheca trichialis, 18 June 1978, J. Hafellner 3899 (GZU!). —Italy: Sudtirol, Mendelgebirge, 
Penegal SW von Bozen, on C chrysocephala, 20 October 1975, J. Hafellner 1095 (GZU!); loc. cit., on 
Chaenotheca sp., 20 October 1975, J. Poelt & J. Hafellner 1094 (GZU!); loc. cit., on Chaenotheca sp., 
20 October 1975,/. Hafellner 1084 (GZU!). 

XIV. MINUTOPHOMA D. Hawksw. gen. nov. 

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata, 
dispersa, globosa vel subglobosa, semi-immersa, nigra, cum muris textura angulari sed inferiore solum 
ex cellulis conidiogenis, ostiolata. Conidiophora desunt. Cellulae conidiogenae enteroblasticae, 
acrogenae, ampulliformes vel breve ampulliformes, phialidicae, non prolifericae, hyalinae sed basi 
plerumque pallide brunneae. Conidia subcylindrica, hyalina, simplicia, laevia, minutissima. 

Conidiomata pycnidial, arising singly, scattered, globose to subglobose, half-immersed, 
black; walls composed of thick-walled dark brown cells forming a textura angularis above, 
these cells readily separating, the lower part consisting only of the conidiogenous cells, the 
upper cells separating to form an irregular ostiole. Conidiophores absent. Conidiogenous 
cells enteroblastic, forming the lower part of the pycnidial wall, acrogenous, ampulliform to 
shortly ampulliform, phialidic, not proliferating, hyaline but with the base often pale brown. 
Conidia not adhering, not catenate, subcylindrical, rounded at the apices, hyaline, simple, 
smooth-walled, minute. 

Type species: Minutophoma chrysophthalmae D. Hawksw. (holotypus). 

Number of species: Monotypic. 

Observations: Minutophoma is most closely allied to Phoma (see p. 49), but differs in the 
extremely small size of the pycnidia and some other important respects; the thick-walled 
angular cells tending to separate around an irregular ostiole, the absence of a differentiated 
wall in the immersed part of the pycnidium, the conidiogenous cells tending to be more 



LICHENICOLOUS COELOMYCETES 



45 







<z> 



lOjLLm 

Fig. 21 Minutophoma chrysophthalmae (IMI 237276 — holotype). A, Vertical section of 
pycnidium. B, Surface view of cells from the upper part of the pycnidium which easily 
separate. C, Conidia. D, Conidiogenous cells. 



elongated and also pigmented below, and the extremely small conidia. An additional feature 
might be the absence of a very deeply pigmented area around the ostiole, but this is not 
emphasized here because the whole of the exposed part of the pycnidium in M. 
chrysophthalmae might be considered as equivalent to this zone. 

The separation of Aster vphoma from Minutophoma is discussed above (p. 8). 

1. Minutophoma chrysophthalmae D. Hawksw. sp. nov. 
(Fig.21A-D) 

Fungus lichenicola. Conidiomata singularia, uniloculata, dispersa, globosa vel subglobosa, semi- 
immersa, nigra, 20-40 pm diam, cum muris textura angulari 5-8 pm latis e cellulis atrobrunneis 
3-4 pm diam sed inferiore solum ex cellulis conidiogenis; ostiolata. Conidiophora desunt. Cellulae 
conidiogenae enteroblasticae, acrogenae, ampulliformes vel breve ampulliformes, phialidicae, non 
prolifericae, hyalinae sed basi plerumque pallide brunneae, 3*5—6 x 2-3*5 //m. Conidia subcylindrica, 
hyalina, simplicia, laevia, 2-3*5 x 1*5 pm. 

Typus: Magna Britannica, Scotia, E. Inverness, Glen Auric, Coille Ruigh na Cuileige, in apotheciis 
Chrysothrix chrysophthalmae (P. James) P. James & Laundon e ligno Pini, 18. vi. 1976, B. J. Coppins 
3196 p.p. (IMI 237276— holotypus!; E— isotypus!). 

Conidiomata pycnidial, semi-immersed, sometimes becoming erumpent and almost 
superficial, scattered over the surface of the host apothecia, arising singly, black, 20^40 pm 
diam; wall in the exposed part of the pycnidium composed of 2-3 layers of cells, 5-8 pm 
thick, dark brown, cells thick-walled, pseudoparenchymatous, tending to be angular, not 
radially compressed, readily separating in squash preparations, 3-4 pm diam, the upper 
cells separating to form an irregular ostiole, the lower part of the pycnidium composed 
almost exclusively of conidiogenous cells, the basal parts of which become slightly pig- 
mented. Conidiogenous cells enteroblastic, forming the lower part of the pycnidial wall, 
arranged in a single layer, acrogenous, ampulliform to shortly ampulliform, phialidic, not 
proliferating, mainly hyaline to subhyaline but with the base often pale brown, 
3*5-6 x 2-3*5 pm. Conidia abundantly produced, subcylindrical, rounded at the apices, not 
adhering, not catenate, hyaline, simple, not distinctly guttulate, smooth-walled, 
2-3*5 x 1-1*5 pm. 

Host: Chrysothrix chrysophthalma (P. James) P. James & Laundon (syn. Micarea 
chrysophthalma P. James), apothecia. Infected apothecia are often richly covered by 



46 D. L. HAWKSWORTH 

pycnidia but otherwise retain their characteristic bright yellow-green appearance and 
continue to produce ascospores. The species consequently has to be interpreted as a 
parasymbiont rather than a parasite. 

Distribution: British Isles (Scotland). 

Observations: A distinctive species which, despite the minute size of the pycnidia, is easily 
detected by their black colour contrasting markedly with that of the host's apothecia. As a 
result of its small size and the minute conidia, Minutophoma chrysophthalmae is unlikely to 
be confused with the other taxa treated in the present revision. 

Additional specimens: British Isles: E. Inverness, Guisachan Forest, south-west of Garve Bridge, on 
Chrysothrix chrysophthalma on Pinus lignum, 26 May 1975, B. J. Coppins et al. 1955 (E!); E. Ross, 
Amat Forest, on Chrysothrix chrysophthalma on Pinus lignum, 28 May 1975, B. J. Coppins (& F. 
Rose 2225 (E!). 

XV. NIGROPUNCTA D. Hawksw. gen. nov. 

Genus lichenicola ad Coelomycetes Sphaeropsidales pertinens. Conidiomata singularia, uniloculata, 
dispersa, subglobosa ad cupuliformia, immersa, plusminusve hyalina, cum muris textura intricata, late 
ostiolata. Conidiophora non vel sparse ramosa, hyalina, septata, e cellulis plusminusve isodiametricis. 
Cellulae conidiogenae holothallicae, acrogenae, integratae, terminales, determinatae, subglobosae ad 
subcylindrices, hyalinae vel ad apicem olivaceae. Conidia catenata, in cirrhum accumulata, 
irregulariter, e cellulis numerosis composita, cellulis subglobosis vel angularibus, atro-olivacea vel 
nigra, plerumque simplicibus, rugulosa. 

Conidiomata pyenidial, arising singly, subglobose at first but becoming cupuliform with 
age, immersed, ± hyaline, largely obscured by the mass of conidia; walls composed of loosely 
interwoven hyaline hyphae forming a textura intricata. Conidiophores forming a compact 
layer lining the inner wall of the pyenidial cavity, not or sparsely branched, hyaline, septate, 
composed of ± isodiametric cells. Conidiogenous cells holothallic, acrogenous, integrated, 
terminal, determinate, subglobose to subcylindrical, hyaline but becoming olivaceous at the 
apex. Conidia catenate at first, accumulating in a cirrhus, irregular, composed of numerous 
subglobose to angular cells, dark olivaceous or black, individual cells mainly simple, 
rough-walled. 

Type species: Nigropuncta rugulosa D. Hawksw. (holotypus). 

Number of species: Monotypic. 

Observations: This genus appears to have no close allies amongst the non-lichenicolous 
fungi. The method of conidiogenesis and the aggregation of cells to form multicellular 
propagules recalls that seen in the lichenicolous hyphomycete genus Sclerococcum Fr. ex Fr. 
(Hawksworth, 1975a, 1979), but there the conidiogenous cells are arranged in superficial 
sporodochia and the conidia are smooth-walled. Amongst the lichenicolous Coelomycetes, 
there is some slight similarity to Vouauxiella uniseptata (see p. 66) but in that taxon the 
conidiogenous cells are much longer and pigmented, the conidia are consistently 1 -septate, 
not aggregating into multi-cellular clumps, have a different type of ornamentation, and are 
produced in a pyenidium with a pigmented and cellular wall. 

The generic name proposed recalls the black-spotted appearance imparted to the host 
thallus. 

1 . Nigropuncta rugulosa D. Hawksw. sp. nov. 

(Figs. 22A-D,23A-C) 

Fungus lichenicola. Conidiomata singularia, uniloculata, dispersa, subglobosa ad cupuliformia, 
immersa. plusminusve hyalina, 100-450 /im lata, cum muris textura intricata 10-20 jum latis, e hyphis 
hyalinis 25-3 //m latis. late ostiolata. Conidiophora non vel sparse ramosa, hyalina, septata, 
10-15 x 2-35 //m, e cellulis isodiametricis. Cellulae conidiogenae holothallicae, acrogenae, integratae, 
terminales. determinatae. subglobosae vel subcylindrices, hyalinae vel ad apicem interdum olivaceae, 



LICHENICOLOUS COELOMYCETES 



47 



L2< J t 




Fig. 22 Nigropuncta rugulosa (IMI 241409 — holotype). A, Infected areolae (x 30). B, Infected 
areolae showing the emergent conidial masses (x 80). C, Vertical section of pycnidium (lower 
part) to show the pycnidial wall and conidiogenous cells (x 1 020). D, Conidia (x 1 020). 



3-4 jum latae, Conidia catenata, in cirrhum accumulata, irregulariter, e cellulis numerosis composita, 
plerumque 20-40 //m diam, atro-olivacea vel nigra, cellulis subglobosis vel angularibus, plerumque 
simplicibus, rugulosis, plerumque 6-8 pirn diam. 

Typus: Austria, Steiermark, Schladminger Tauren, Kleinsolk-Obertal, west. Abhange der Kessel- 
spitze, alt. 1700 m, in thallo lichenis ignoti e saxis schistosis, 12. viii. 1974, ./. Hafellner 3929 (IMI 
241409 — holotypus!; herb. Hafellner — isotypus!). 

Conidiomata pycnidial, arising singly, immersed in areolae of the host, the adjacent 
thalline tissue sometimes proliferating to form a distinct raised margin around the spore 
mass which recalls a thalline exciple, only exceptionally more that one pycnidium in each 
areole, ± hyaline but black when viewed from above as a result of the dense spore mass, 
100-450 [im wide; ostiolate, the ostiole ± equal to the width of the pycnidium in older 
conidiomata, mainly ± circular but the margin sometimes invaginated or angular, not 
clearly differentiated but forming by an irregular opening caused by an expansion of the 
pycnidium, the pycnidia are at first subglobose or flask-shaped but become cupulate to 
almost acervular with age; walls poorly demarcated from the host tissues, mainly hyaline but 
sometimes becoming olive-tinged, especially towards the upper parts, 10-20//m thick, the 



48 



D. L. HAWKSWORTH 





lOjim 

Fig. 23 Nigropuncta rugulosa (IMI 241409 — holotype). A, Conidiogenous cells and detail 
of pycnidial wall. B, Conidiogenous chain with developing conidia. C, Cells squashed 
from the irregular multicellular conidia. 



outer part composed of 2-5 irregular layers of loosely to moderately compacted interwoven 
hyphae, hyphae thick-walled, hyaline, 2*5-3 jum wide, inner part comprising thinner-walled 
mainly isodiametric loosely packed pseudoparenchymatous cells, 2—3*5 jum diam, these 
scarcely distinguishable from conidiophores. Conidiophores forming a compact layer lining 
the inner wall of the pycnidial cavity, simple or sparsely branched, arranged ± parallel to 
one another, hyaline, septate, 10-15 x 2—3*5 //m, composed of almost isodiametric cells, 
somewhat constricted at the septa. Conidiogenous cells holothallic, integrated, terminal, 
determinate, subglobose to subcylindrical, resembling the conidiophore cells but sometimes 
becoming olivaceous at the apex, 3^ jum wide. Conidia arising in short basipetal chains, 
tightly adhering to those produced from adjacent conidiogenous cells to form multicellular 
propagules, these very irregular in shape but mainly 20^0 jum overall, deep olive to almost 
black, extruded through the ostiolar opening as a dense black cirrhus which is clearly seen 
with a x 10 lens, individual cells mainly simple, thick- walled, subglobose to angular through 
compression by adjacent cells, walls roughened with a very irregular granular-lacerate 
ornamentation, this perhaps originating from exfoliating layers of wall tissue, cells mainly 
6-8 //m diam. 

Host: On an unidentified sterile grey areolate crustose lichen thallus. The areolae are not 
discoloured in any way and algal cells close to the pycnidia retain their usual colour also. 
The species is evidently parasymbiotic rather than parasitic, but could even be lichenized 
(see below). 

Distribution: Austria. Known only from the type collection with certainty, but perhaps 
also present in Germany (see below). 

Observations: This is a most distinctive species macroscopically and superficially recalling 
Thelomma siliceum (Fee) Tibell (see Tibell, 1976: figs. 15-16); however, microscopic 
examination soon reveals that it is not an ascomycete. 

Within the host thallus, it seem impossible to distinguish hyphae of the pycnidium from 



LICHENICOLOUS COELOMYCETES 49 

those forming the tissues of the host. Further, most areolae have only a single pycnidium and 
the pycnidia are evenly distributed over the material, and not in any way localized. These 
facts, together with the consideration that the thallus and its algae are maintained in a 
healthy condition, makes it necessary to speculate whether the present fungus is actually an 
independent lichenized coelomycete. In the absence of additional material for study and 
detailed ultrastructural investigations, it seems impossible to resolve this question at the 
present time. 

One factor which may support the parasymbiotic thesis is a report of what may be the 
same fungus on Lecanora alphoplaca (Wahlenb.) Ach. by Gerber (1931 : 481 figs. 14-15). 
This author illustrated an unnamed fungus on this host collected by Zopf at 'Oberwinkel in 
Groden' which has some important similarities to Nigropuncta rugulosa, particularly in the 
macroscopic appearance of the infection and the formation of dark-coloured multicellular 
conidia which were very irregular in shape and composed of subglobose to angular cells. 
Interestingly, he appears to have considered the structure referred to above called the 
pycnidial wall as the invaginated cortical layer of the host. I have not been able to locate the 
material Gerber studied, and from his paper alone cannot confidently assert that he had 
the same taxon because sufficient information on dimensions and conidium formation was 
not provided. If the two are conspecific, however, as Gerber had a host he could determine, 
then N. rugulosa must be considered a parasymbiont and not an independent lichen. 

The dark olive colour of the conidia is particularly characteristic and must be due to a 
pigment (or pigments) different from those present in other dark-spored lichenicolous 
Coelomycetes. The lacerate-granulose roughening of the conidia is also an unusual feature. 
This may conceivably originate either from an exfoliation of the conidial wall layers, or be 
wall remnants produced during conidiogenesis. The pigmentation precludes a more detailed 
light microscopic study, and ultrastructural investigations are needed to firmly determine the 
nature of this roughening and any relationship it might have to the process of conidiogenesis. 

XVI. PHOMA Sacc. nom. cons. 

Michelia 2: 4 (1880). 

See Sutton (1980 : 378) for generic synonyms. 

Conidiomata pycnidial, arising singly or aggregated, globose, immersed or partly 
erumpent, brown to dark brown, ostiolate; walls usually thin, composed of a few layers (often 
1-3) of cells, cells pseudoparenchymatous, pale brown to brown, more deeply pigmented 
around the ostiole in many species, rather thin-walled, textura angularis. Conidiophores 
normally absent. Conidiogenous cells enteroblastic, lining the pycnidial cavity, short- 
cylindrical to doliiform, phialidic, not proliferating, hyaline. Conidia arising singly, not 
catenate, variable in shape in different species, ellipsoid, subcylindrical, fusiform, pyriform 
or globose, rounded at the apex and base or with the base slightly truncated, hyaline, simple 
when mature (exceptionally becoming 1 -septate), sometimes distinctly guttulate, thin- 
walled, smooth-walled. 

Type species: Phoma herbarum Westend. 

Teleomorph (perfect states): A wide range of loculoascomycetes are described as having 
anamorphs referrable to Phoma, including species of Leptosphaeria Ces. & de Not. and 
Pleospora Rabenh. ex Ces. & de Not. Most Phoma species, however, remain unconnected to 
teleomorphs. 

Number of species: According to Sutton (loc. cit.) many more than 2000 species have 
been described in this genus, which was traditionally employed for all caulinicolous 
Coelomycetes with small, hyaline, simple conidia. The genus is now employed for species 
from many habitats (and includes taxa from a very wide range of substrates) and perhaps 
really should only include about 50 species; a key to 27 is included in Sutton (loc. cit.). Five 
lichenicolous species are now known. 



50 



D. L. HAWKSWORTH 




Fig. 24 Phoma caloplacae (UPS — holotype). A, Surface view of pycnidium. B, Surface 
view of pycnidial wall. C, Conidiogenous cells and pycnidial wall. D, Conidia. 

Observations: Phoma-like fungi on lichens have often been placed in Phyllosticta Pers. ex 
Desm., a genus formerly adopted for similar fungi on leaves as opposed to stems, presumably 
in the belief that lichen thalli approximated more closely to 'leaves' than 'stems'. The type 
species of Phyllosticta, however, is in any case very different, conforming to the concept of 
Phyllostictina Sydow, and can not be used for such fungi (Punithalingam, 1974 : 60-61). The 
lichenicolous species referred to Phoma here agree closely with the concept of Phoma 
adopted by Sutton (loc. cit.). 

Modern work on species concepts in Phoma has emphasized studies of their behaviour in 
pure culture and it would obviously be of interest to obtain the lichenicolous species in 
culture and then compare the isolates with the known non-lichenicolous taxa. 



Key to the lichenicolous species 

1 Conidia broadly to narrowly ellipsoid 2 

Conidia subglobose, (4-)5-6(-7) //m diam; on Caloplaca cerina . • Phoma caloplacae (p. 50) 

2(1) Conidia narrowly ellipsoid, usually less than 2*5 //m wide 3 

Conidia broadly ellipsoid, 4-5 - 5(-6)x 2-5-3-5(-4)/^m; on Physcia aipolia and Physconia 
pulverulacea Phoma physciicola (p. 56) 

3(2) Conidia to 2 //m wide 4 

Conidia (4-)4'5-6(-7) x 2-2*5(-3) //m; on Peltigera malacea • . . Phoma peltigerae (p. 54) 

4(3) Conidia 5-7 x 1-5-2 //m; on Parmelia species Phoma cytospora (p. 5 1 ) 

Conidia 35-5 x 15-2 //m; on an Usnea species Phoma dubia (p. 53) 

1. Phoma caloplacae D. Hawksw. sp. nov. 
(Fig. 24A-D) 

Fungus lichenicola. Conidiomata singularia, uniloculata, dispersa, subglobosa, immersa sed 
erumpescentia, atrobrunnea, (50-)70-100(-130)/im lata, cum muris textura angulari usque 10-15 //m 



LICHENICOLOUS COELOMYCETES 5 1 

latis, e cellulis 5-8 x 3-4 jum. Cellulae conidiogenae enteroblasticae, acrogenae, subglobosae vel late 
obpyriformes, phialidicae, non prolifericae, hyalinae, 5-6 jum diam. Conidia subglobosa, hyalina, 
simplicia, laevia, (4-)5-6(-7) p.m diam. 

Typus: U.S.S.R., Guv. Jenisejsk, Stolba, lat. bor. 60°20 , N, in apotheciis Caloplacae cerinae (Ehrh. ex 
Hedw.)Th. Fr., l.vii. 1876, M. Brenner 1027b p.p. (UPS— hqlotypus!). 

Conidiomata pycnidial, immersed at first, the upper \ erumpent at maturity, arising 
singly, scattered, dark brown, subglobose, (50-)70-100(-130)//m diam, ostiolate, cells 
surrounding the ostiole dark brown, darker than the remaining wall which is subhyaline to 
pale brown; wall 3-5 layers of cells thick, 10-15 pm thick, cells pseudoparenchymatous, the 
outermost brown to dark brown with the walls thickened, the inner subhyaline with thinner 
walls, generally somewhat radially compressed and angular, 5-8 pm diam in surface view 
and mainly 3-4 pm thick in section. Conidiogenous cells arising from the inner wall of the 
pycnidium, lining the pycnidial cavity, subglobose to broadly obpyriform, hyaline, 
smooth-walled, phialidic, not proliferating, 5-6 pm diam. Conidia abundant, subglobose 
but often somewhat angular due to mutual compression within the pycnidial cavity, hyaline, 
simple, sometimes guttulate, smooth-walled, (4-)5-6(-7) /im diam. 

Host: Caloplaca cerina, apothecia. Infected discs become discoloured dark brown to 
blackish and ascospore production is inhibited by the presence of the fungus. 

Distribution: U.S.S.R. Known only from the type collection. 

Observations: Readily separated from the other species accepted in the genus by the 
subglobose to angular, and not ellipsoid, conidia. 

2. Phoma cytospora (Vouaux) D. Hawksw., Trans. Br. mycol. Soc. 67 : 56 (1976). 
(Fig. 25A-C) 

Phyllosticta cytospora Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 193 (1914). 

Type: France, Blainville, on Parmelia caperata, 4 March 1906, M. Missie (herb. Vouaux — 
neotype!). 

Icones: Hawksworth & Punithalingam, Trans. Br. mycol. Soc. 61 : 62 fig. 4 (1973). 

Conidiomata pycnidial, immersed, arising singly or more commonly in groups, scattered, 
black, subglobose, 40-80 pm diam, ostiolate; wall 2-3 layers of cells thick, mainly 
7-10 //m thick but to about 15 pm near the ostiole, brown to dark brown, cells 
pseudoparenchymatous, not markedly radially compressed, 5-7 pm long and 3^4 /im tall, 
the inner cells subhyaline to pale brown and with somewhat thinner walls. Conidiogenous 
cells arising from the inner wall of the pycnidium, lining the pycnidial cavity, subglobose to 
broadly obpyriform, hyaline, smooth-walled, phialidic, not proliferating, 2-3 x \-2 pm. 
Conidia abundant, narrowly ellipsoid to slightly curved, one apex slightly truncated, 
hyaline, simple, sometimes 1-2 guttulate, smooth-walled, 5-7 x F5-2 pm. 

Hosts: Parmelia caperata (L.) Ach., P. laevigata (Sm.) Ach., P. perlata (Huds.) Ach., P. 
reticulata Taylor, and P. sulcata Taylor, thalli. Necrotic decolourized patches \-^ mm diam 
are formed, each of which includes 5-18 pycnidia, the boundary with healthy tissue being 
marked by a blackened zone of tissue 100-250 //m wide; the patches are usually discrete 
and damage localized but occasionally they may coalesce, particularly on P. reticulata, 
killing large areas of the thallus. Reported by Vouaux (1914) and Keissler (1930) only on P. 
caperata. 

Distribution: British Isles, France, and possibly Germany (see p. 83). 

Observations: The decolourized necrotic patches produced by this species recall those 
formed on the same or allied hosts by Cornutispora lichenicola and Lichenoconium erodens. 

Phyllosticta cytospora was originally described from material of Parmelia caperata 
collected in the Toret de Vitrimont en Meurthe-et-Moselle' by Vouaux. This material is no 
longer present in Vouaux's herbarium (Rondon, 1970) but there is a further collection made 




Fig. 25 Phoma cytospora (IMI 164975). A, Vertical section of pycnidium. B, Conidio- 
genous cells and pycnidial wall. C, Conidia. Reproduced from Hawksworth & Punithalingam 
(1973:62). 



LICHENICOLOUS COELOMYCETES 



53 







25/im 




Fig. 26 Phoma dubia (E — holotype). A, Vertical section of pycnidium. B, Conidiogenous 
cells and pycnidial wall. C, Conidiogenous cells. D, Conidia. 



at Blainville in 1906 labelled 'Phyllosticta cytospora mini' present which agrees in all details 
with Vouaux's original description and later collections; the Blainville specimen is 
consequently designated as neotype for this name here. 

The name Phyllosticta lichenicola Allescher may have been partly based on a fungus very 
close to Phoma cytospora (see p. 83). Phoma caperatae Vouaux, however, seems unlikely 
to represent P. cytospora (see p. 80). 

Additional specimens: British Isles: England, S. Devon, Slapton Ley Nature Reserve, on Parmelia 
perlata on Quercus, 30 March 1972, D. L. Hawksworth 2695 (IMI 164975!), he. eit., on P. reticulata on 
Acer pseudoplatanus, 25 August 1978, D. L. Hawksworth 4112 (IMI 231655!), loc. cit., on P. sulcata, 25 
August 1976, D. L. Hawksworth 4322 (IMI 206364!); S. Devon, Killerton Park, on P. reticulata on 
Quercus, 3 September 1978, D. L. Hawksworth 4849 (IMI 231732!); S. Devon, Start Point, nr 
Langerstone Point, on P. perlata on Salix, 27 August 1972, D. L. Hawksworth 3075 (IMI 168590!); S. 
Devon, Cornwood, Dendles Wood, on P. laevigata, 29 February 1976, D. L. Hawksworth 4293a (IMI 
201661a). Scotland, Dumfriesshire, nr Gretna, on P. sulcata on Ulmus, 6 August 1972, B. J. Coppins 
(E!); Argyll, S side Loch Sunart, on P. laevigata, 14 August 1972, S. R. Davey (IMI 250032/7/ 

3. Phoma dubia (Lindsay) Sacc. & A. Trotter, in Saccardo, Syll. Fung. 22 : 897 ( 1 9 1 3)*. 
(Fig. 26A-D) 



*Phoma dubia Rupprecht (Sydowia 11 : 128, 1958), described from Stellated graminea in Hungary, is a later 
homonym which must be rejected under Art. 64. 1 . 



54 D. L. HAWKSWORTH 

Phymatopsisdubia Lindsay, Trans. R. Soc. Edinb. 24 : 442 (1866). 
Type: New Zealand, Otago, Pelichet Bay, in the bush, on Usnea sp. (apothecia), 17 December 
1861, W. L. Lindsay (E— holotype!). 

Icones: Lindsay, Trans. R. Soc. Edinb. 24 : pi. 30 figs. 36^1 (1866). 

Conidiomata pycnidial, immersed in the hymenium of the host, only the ostiole 
sometimes slightly protruding above the level of the epithecium, scattered to loosely 
aggregated, sometimes almost confluent, subglobose, 25-60 jum diam, ostiolate; wall mainly 
1-3 layers of cells thick, mostly 3*5—5 jum thick but to 8 jum near the ostiole, cells 
pseudoparenchymatous, subglobose to somewhat polyhedral, subhyaline except in the 
vicinity of the ostiole and there thicker-walled and golden-brown, mainly 2-3 jum diam. 
Conidiogenous cells arising from the inner wall of the pycnidium, short-ampulliform to 
obpyriform, hyaline, smooth-walled, phialidic, not proliferating, 4-8 x 3-5 jum. Conidia 
abundant, narrowly ellipsoid to almost bacilliform, sometimes slightly truncated at the base, 
hyaline, simple, 0-2 guttulate, smooth-walled, 3*5 x L5-2 //m. 

Host: Usnea sp., apothecia. The host was originally named as U. barbata var.florida (L.) 
Fr. (i.e. U.florida (L.) Wigg.) by Lindsay but I am not sure that it really belongs there; certain 
identification must await a re-evaluation of that genus in New Zealand. The infected parts of 
the apothecial disc become discoloured and brownish suggesting that Phoma dubia is a mild 
pathogen. 

Distribution: New Zealand. Known only from the original collection. 

Observations: This fungus is clearly separated from the other lichenicolous Phoma species 
treated here by the minute conidia. 

4. Phoma peltigerae (P. Karsten) D. Hawksw., Trans. Br. mycol. Soc. 74 : 381 (1980). 
(Fig.27A-E) 

Phyllosticta peltigerae P. Karsten, Hedwigia 23 : 62 (1884). 
Type: Finland, Tavastia australis, Tammela, Kyto, on Peltigera malacea, 4 May 1869, P. A. 
Karsten (H-KARST 2040— holotype!). 

Conidiomata pycnidial, ± completely immersed at first, the upper \ erumpent at 
maturity, arising singly, scattered, black, subglobose, (75-)100-150(-200)/im diam, 
ostiolate, cells surrounding the ostiole dark brown to black and darker than the remainder of 
the wall; wall 1-2 layers of cells thick, 5-8 p,m thick, cells pseudoparenchymatous, the 
outermost dark brown with the external wall thickened, not markedly radially compressed, 
5-9 /urn long and 4-7 /im tall, the inner cells subhyaline to pale brown and with thinner 
walls. Conidiogenous cells arising from the inner wall of the pycnidium, lining the pycnidial 
cavity, subglobose to broadly obpyriform, hyaline, smooth-walled, phialidic, not pro- 
liferating, 4-6 jum diam. Conidia abundant, narrowly ellipsoid, rounded at the apices, 
hyaline, simple, sometimes guttulate, smooth-walled, (4-)4'5-6(-7) x 2-2 5(-3) jum. 

Host: Peltigera malacea (Ach.) Funck, thallus. The host in the type collection was 
originally determined as P. canina (L.) Willd., and this has been given as the host in all 
previous references to this species. On P. malacea the infected area becomes entirely 
decolourized, the whitened patches being 5-10 mm across and very sharply delimited from 
the healthy tissues; a slight blackening of the host occurs just at the margin of the infection 
spot. Phoma peltigerae consequently appears to be pathogenic to this species. This fungus 
has also been reported from P. horizontalis (Huds.) Baumg., P. polydactyla (Necker) Hoffm. 
and P. rufescens (Weis) Humb. by Vouaux (1914 : 193), and Keissler(1930 : 536) mentions a 
report on Platismatia glauca (L.) Culb. & C. Culb.; these four host records require 
substantiation. 

Distribution: Finland. Known to me only from the type collection but mentioned as 
present in France by Bouly de Lesdain (1910:276) and Vouaux (1914: 193), and in 
Germany by Keissler(1930 : 536, 1933 : 392). 



LICHENICOLOUS COELOMYCETES 

B 



55 





E ^§1% 

1%^ y 




10/im 

Fig. 27 Phoma peltigerae (H-KARST 2041 — holotype). A, Surface view of pycnidium. B, 
Vertical section of pycnidium. C, Surface view of pycnidial wall. D, Conidiogenous 
cells and pycnidial wall. E, Conidia. 



Observations: Phoma peltigerae recalls P. cytospora but differs in the broader conidia, 
thickening of the pycnidial wall cells, and larger pycnidia, as well as in occurring on quite 
different hosts. 

Phyllosticta peltigerae was originally described from 'Mustiala' but no material under this 
name from that locality could be found in H-KARST (O. Vitikainen, in litt.). However, in a 
packet incorrectly labelled as 'Stagonopsis peltigerae' in W. Nyberg's hand (an amateur 
mycologist who once helped in H) was a slip of paper with the herbarium name 'Phoma 
peltigerae' in Karsten's writing, conidial characters and notes conforming to the published 
description of Phyllosticta peltigerae, and the locality name Kyto. This is a place name in 
Tammela parish, situated west of Mustiala (O. Vitikainen, in litt.), and it appears that when 
publishing this fungus Karsten (a) used a nearby better-known locality name, and (b) as 
Phoma and Phyllosticta were then separated according to whether the fungus was on a stem 
or leaf, decided that the thallus of Peltigera was leaf-like not stem-like. H-KARST 2040 is 
consequently considered to be the holotype of Phyllosticta peltigerae. 



56 



D. L. HAWKSWORTH 




10/im 

Fig. 28 Phoma physciicola (W 1910/609 — holotype). A, Vertical section of pycnidium. B, 
Conidiogenous cells and pycnidial wall. C, Surface view of pycnidial wall. D, Conidi- 
genous cells. E, Conidia. 



5. Phoma physciicola Keissler, Hedwigia 50 : 294 (1911). 
(Fig. 28A-E) 

Type: Austria, Steiermark, Gams bei Hieflau, on Phvscia aipolia, June 1910, K. von Keissler (W 
1910/609— holotype!). 

Icones: Keissler, Hedwigia 50:295 fig. la-c (1911). — Keissler, Rabenh. Krvpt.-Fl. 8:539 figs. 
102-103(1930). 

Conidiomata pycnidial, immersed at first, with the upper \ erumpent at maturity, arising 
singly or in small groups, black, subglobose, 100-150 jum diam, ostiolate, cells surrounding 
the ostiole dark brown to black and darker than the remainder of the wall; wall 1-3 layers of 
cells thick, mainly 7-10 //m thick but to 15 jum near the ostiole, cells pseudoparenchy- 
matous, the outermost brown with moderately thickened walls, not markedly radially com- 
pressed, 5-7 p,m diam in side view, 5-10 //m diam and rather angular in surface view, the 
inner cells pale brown to subhyaline and with thinner walls. Conidiogenous cells arising 
from the inner wall of the pycnidium, lining the pycnidial cavity, subglobose, hyaline, 
smooth-walled, phialidic, very rarely proliferating, 4-7 jum diam. Conidia abundant, 
broadly ellipsoid, rounded at the apices, hyaline, simple, frequently with 2 or more guttules, 
smooth-walled, 4-5'5(-6) x 2-5-3'5(-4) jum. 

Host: Physcia aipolia (Humb.) Fiirnrohr and Physconia pulverulacea Moberg, apothecia. 
Also cited by Keissler (1930:542) from Baeomyces rufus (Huds.) Rebent. (thallus), 
Parmelia caperata (L.) Ach. (thallus) and P. glabratula (Lamy) Nyl. (thallus) but these 
reports are dubious and require confirmation. In the holotype, as many as 15 pycnidia may 
occur in a single apothecium of the host in which they are rather strictly confined to the 
thecium, scarcely extending down into the hypothecium. Infected apothecia retain their 
black colour but lose their characteristic whitish pruina; ascospores still formed in the most 
heavily infected apothecia studied. 



LICHENICOLOUS COELOMYCETES 57 

Distribution: Austria and Sweden. 

Observations: Phoma physciicola is separated from other species of the genus accepted 
here by the shape and size of the conidia. Phoma caperatae Vouaux was treated first as a 
variety of P. physciicola and later subsumed under this species by Keissler (1930 : 542), but it 
is most unlikely that these taxa are conspecific (see p. 80). 

Additional specimens: Sweden: Smaland, Aneboda s:n, Aneboda limnologiska laboratorium, pa 
lonn, i alle, on Physconia pulxerulacea, 27 July 1947, R. Santesson (UPS!); Sodermanland, Bankyska 
s:n, Sturehav, alle trad, on P. pulverulacea, 2 1 July 1944, R. Santesson (UPS!). 

XVII. PSEUDOSEPTORIA Speg. 

An. Mus. nac. B. Aires 20 : 388 (1910). 
Lunospora Frandsen, Meddr Kgl. Veter. Landboh. Kobenh. 26 : 70 (1943). 

Conidiomata pycnidial, arising singly or aggregated into short rows, globose, immersed to 
erumpent, brown to black, ostiolate; walls composed of a few layers of cells, cells 
pseudoparenchymatous, thin- to moderately thick-walled, pale to greenish or dark brown, 
textura angularis. Conidiophores absent. Conidiogenous cells holoblastic, lining the inner 
wall of the pycnidial cavity, acrogenous or sympodial, subcylindrical to ampulliform, often 
proliferating, then with distinct annellations, hyaline. Conidia not catenate, falcate, fusiform 
or cymbiform, attenuated at the apex, attenuated to truncate at the base, hyaline, simple, 
smooth-walled. 

Type species: Pseudoseptoria donacis (Pass.) B. Sutton (syn. P. donacicola Speg.). 

Number of species: Four species are treated by Sutton (1980), all of which are 
graminicolous. A fifth lichenicolous species is added to the genus here. 

Observations: The genus Pseudoseptoria has been revived by Sutton (1980) for several 
graminicolous species hitherto referred to Selenophoma Maire, a genus that differs in both 
pycnidium structure and the method of conidiogenesis. The inclusion of Phoma usneae 
Vouaux in the genus expands the generic concept slightly to embrace species in which the 
conidia are not strongly falcate. It is probable that some additional non-lichenicolous and 
non-graminicolous will eventually be found to require transfer here. 

1. Pseudoseptoria usneae (Vouaux) D. Hawksw. comb. nov. 
(Fig. 29A-B) 

Phoma usneae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 196 (1914). 

Type: Italy, Tirol, Bolzano [Bolzen], Mendola [Mendel], wood by Mt Roen [Roen], on Usnea 
fdipendula aggr. (thallus), 13 August 1896, F. G. C. Arnold [Lich. Exs. no. 1718 p.p.] 
(K — isotype!). 

Exsiccatae: Arnold, Lich. Exs. no. 1718 p.p. (K!; sub Epicoccum usneae Anzi). 

Conidiomata pycnidial, immersed at first but becoming erumpent with age, arising singly, 
scattered, black, subglobose, ostiolate, 75-100(-200)//m diam when mature; wall about 3-4 
layers of cells thick, cells moderately thick-walled, pseudoparenchymatous, forming a 
textura angularis, subglobose to polyhedral, greenish brown, 4-7 pm diam. Conidiogenous 
cells holoblastic, lining the inner wall of the pycnidial cavity, subcylindrical to ampulli- 
form, percurrently proliferating, to 4 annellations seen, hyaline, smooth-walled, rarely 
proliferating sympodially, 6-12 x 3-5 pm. Conidia cymbiform, tapering to the rounded 
apex but with an abruptly truncated base, base 2-3 jum wide, hyaline, smooth-walled, 
simple, often with numerous small guttules, ( 1 0—) 1 2— 1 4(— 16) x (3-)3-5^1//m. 

Host: Usnea fdipendula Stirton aggr., thallus. The fungus is present on some of the smaller 
branches but very few pycnidia were found on the isotype studied. Infection, at least in its 
early stages, does not appear to damage the host significantly. The apothecia in the type 



58 



D. L. HAWKSWORTH 

B 






Fig. 29 Pseudoseptoria usneae (K — isotype). A, Conidiogenous cells. B, Conidia. C, Surface view 

of cells of pycnidial wall. 



collection are infected with Lichenoconium usneae (Anzi) D. Hawksw., and some dark 
brown torulose mycelium and galls that may be young Abrothallus usneae Rabenh. are 
present on other parts of the thallus. 

Distribution: Italy and Spain (fide Santesson, 1960 : 519). 

Observations: Vouaux described this taxon on the basis of the example of Arnold's 
exsiccatum in Harmand's herbarium. This could not be located amongst Harmand's 
herbarium in Angers (M. Guerlesquin, in litt.) but one collection of this number in K was 
found to have a few pycnidia producing conidia conforming so precisely to Vouaux's 
description that there can be no doubt they belong to the same fungus. The only 
discrepancies noted were that Vouaux indicated the pycnidia were 100-200 //m diam and 
that he gave the conidiogenous cells as only 3-6 x 2 //m. So few pycnidia were present on 
the specimen in K that no microtome sections were prepared; a more detailed account of the 
pycnidium structure must consequently await the availability of further material of this 
taxon. 

This species occupies a rather isolated position in Pseudoseptoria as currently 
circumscribed in view of the relatively broad and ± straight, rather than curved or falcate, 
conidia. Such a difference scarcely merits a generic separation, however, as it seem to agree 
with the type species of the genus in all other details. 



LICHENICOLOUS COELOMYCETES 59 

Keissler (1960 : 416) indicated that this fungus could be distinguished from the normal 
pycnidia of Usnea species in that the conidiogenous cells were simple, but there are many 
other differences also. For example, the conidia of Usnea are, contrary to some earlier 
reports, bacillariform with a marked swelling just above the base so that they appear 
pendulum-shaped (Swinscow & Krog, 1976 : 264-5). 

XVIII. PYRENOTRICHUM Mont. 

Annls Sci. nat. (Bot.) II 20 : 376 (1 843). 

Chlorocyphella Speg., An. Mus. nac. B. Aires 19 : 279 (1909). 

Conidiomata ('campylidia', 'orthidia') helmet-shaped, erect, convoluted or becoming 
adpressed, superficial, arising singly, not aggregated into stromata, pale fawn to shades of 
blue-green or blue-black, wall thick at the base but tapering towards the tip, outer convex 
surface composed of arachnoid hyphae, central tissue and inner concave surface of hyaline to 
pigmented thick-walled pseudoparenchymatous cells, textura angularis. Conidiophores 
mainly in the lower part of the concave surface, simple or branched near the base, septate or 
unicellular, conidiogenous cells arising at ± the same level, ellipsoid to subcylindrical. 
Conidiogenous cells probably enteroblastic, phialidic, not proliferating, lacking distinct 
collarettes, hyaline, smooth-walled, subcylindrical. Conidia arising singly, ellipsoid, 
pyriform, clavate, filiform or scolecosporous, sometimes variously curved, O-multiseptate, 
not or distinctly branched, hyaline, smooth-walled. 

Type species: Pyrenotrichum splitgerberi Mont. 

Number of species: Three species have validly published names in Pyrenotrichum, and 
Santesson (1952 : 41) recognized six additional species which have not yet been formally 
published. All these nine species are parasymbionts on tropical foliicolous lichens and it is 
probable that there are additional species awaiting description on other crustose lichens on 
bark in the tropics (see below). 

Observations: Pyrenotrichum species are evidently widespread in the tropics and have 
been studied in some detail by Santesson; unfortunately only a brief account of his important 
investigations has so far appeared (Santesson, 1952:40^11). I have not attempted to 
duplicate his research and consequently describe in full only the type species of the genus 
and note one of the other species with a validly published name represented in IMI. In 
addition to these two species, P.foliicola mentioned under P. staurosporum, and P.filiferum 
mentioned under P. splitgerberi, Santesson accepts five species with simple or 1 -septate 
conidia; ellipsoid conidia occur in P. bicolor R. Sant. ined. (on Lopadium flammeum Mull. 
Arg.), P. mirum R. Sant. ined. (on L. tayabasense (Vainio) Zahlbr.), and P. atrocyaneum R. 
Sant. ined. (on Lopadium sp.); and ± pyriform to ± clavate conidia in P. irregulare R. Sant. 
ined. (on four Sporopodium species), and P. podosphaera R. Sant. ined (on Sporopodium 
sp.). 

The genus is evidently not restricted to foliicolous lichens as I have seen two collections 
referrable to it on sterile crustose lichens on bark from the tropics (Ecuador: Galapagos 
Island, Isla Pinta, near volcano in south-central part, on Tournefortia sp., 8 July 1976, H. 
Sipman, COLO-L63608, IMI 214571!; St Helena: Ascension Island, Green Mountain, nr 
summit, on Podocarpus sp. bark, 30 October 1976, P. W. James, IMI 251280!). Whether 
these represent additional new species is unclear, resultant upon the publication of 
Santesson's revision. 

It is possible that some of the allegedly lichenized foliicolous Coelomycetes described by 
Brazilian workers in recent years also belong to Pyrenotrichum as interpreted by Santesson, 
for example Acleistomyces rionegrensis Bat. & Maia with ellipsoid, simple, conidia (Batista, 
1961). The status of many of the taxa introduced by these workers requires a critical 
re-evaluation; the generic names involved are compiled by Vobis & Hawksworth (1981). 

Keissler (1927: 159) listed generic names 'Campylidium Mull. Arg.' and 'Orthidium 
Mull. Arg.' as synonyms of Chlorocyphella: this is quite incorrect as Muller (1881 : 111, 



60 D. L. HAWKSWORTH 

1 890 : 202) introduced these as anatomical terms for conidiomata of different shapes and not 
as generic names. Neither of these names is validly published as a genus. 

I found the method of conidiogenesis in P. splitgerberi difficult to ascertain conclusively, 
even by differential interference contrast. Although the conidia are often large, at the point of 
secession a structure only about 1*5 jum wide must be resolved; ultrastructural studies will 
be necessary to conclusively determine this matter. 

Key to the species treated 

1 Conidia unbranched, filiform, multi-septate, arcuate to C- or S-shaped or variously 

convoluted, 55-80 x 2-3 jum Pyrenotrichum splitgerberi (p. 60) 

Conidia with a main stem and 2-4 branches arising at a single locus about halfway along the 
conidum, arms at acute angles, straight to slightly flexous, 60-85 x 2*5-3-5 /im 

Pyrenotrichum staurosporum (p. 63) 

1. Pyrenotrichum splitgerberi Mont., Annls Sci. nat. (Bot.) II, 20 : 377 (1843). 
(Figs. 30A-F,31A-B) 

Lccicka irregularis Fee, Bull. Soc. bot. Fr. 20 : 318 (1873). 

Melophia woodsiana Sacc. & Berl., in Saccardo, Svli Fung. 3 : 659 (1884)*. 

Cyphella subcyanea Ell. & Ev., J. Mycol. 2 : 37 (1 886). 

Cyphella aeruginascens P. Karsten, Hedwigia 28 : 191 (1889). 

Chlorocyphella aeruginascens (P. Karsten) Keissler, Annln naturh. Mus. Wien 41 : 1 59 (1927). 

? Trichosperma cyphelloidea Hohnel, Sber. Akad. Wiss. Wien 1, 116 : 145 (1907)*. 

?Trichosperma aeruginosa Hohnel, Sber. Akad. Wiss. Wien 1, 118 : 1 530 (1909)*. 

Chlorocyphella subtropica Speg., An. Mus. nac. B. Aires 19 : 279 (1909). 

Chlorocyphella aeruginascens var. convoluta Keissler, Annln naturh. Mus. Wien 41 : 161 (1927). 

Chlorocyphella lichenicola Keissler, in Zahlbruckner et al, Trans. Proc. N.Z. Inst. 59 : 3 13 (1928). 

Icones: Keissler, Annln naturh. Mus. Wien 41 : 161 fig. \a-b (1927). — Mameli-Calvino, Nuovo G. 
bot. ital. II, 37 : pi. 19 figs. 1-16 (1930).— Montagne, Annls Sci. nat. (Bot.) II, 20 : pi. 16 fig. 
2a-m (1843).— Rizzini, Archos Jard. bot., Rio deJ. 12 : figs. 1-7 (1952). 

Conidiomata helmet-shaped, ± erect, convoluted, or geniculately adpressed, entirely 
superficial, almost fawn to blue-green or blue-black, mat, arising singly, scattered, often 
rather evenly and sparsely distributed over the host thallus, very variable in size and shape, 
mainly 300-800 jum long and 250-600 jum wide when mature; wall 40-60 //m thick at the 
base but tapering to about 10 jum thick at the tip, in vertical section seen to be 
composed of two layers, an irregular outer layer of arachnoidly branched thick-walled 
hyphae 3-3 • 5 jum wide with granular incrustations often evident on their walls, and air- 
spaces present in the gaps formed by the hyphae, inner layer of thick-walled pseudoparen- 
chymatous cells, textura angularis, cells irregularly subglobose to ellipsoid or polyhedral, 
mainly 4-9 jum diam, without air spaces, subhyaline to olivaceous, the cells towards the 
inner concave side adjacent to the conidiogenous layer most deeply pigmented, those above 
the conidiogenous region often with papilliform projections. Conidiophores simple or 
branched near the base, septate or unicellular, hyaline, ellipsoid to subcylindrical, with 1-3 
conidiogenous cells arising at about the same level, to 10x3 //m, restricted to the concave 
surface of the conidiomata and not extending more than about \ of the length of the 
conidiomata in the specimens sectioned. Conidiogenous cells probably enteroblastic, 
phialidic with minute apical thickenings, arising from the conidiophores or sometimes 
apparently directly from the innermost wall cells, hyaline, smooth-walled, subcylindrical, 
slightly wider at the base but not ampulliform, mainly 10-12 x2'5-3'5(-4)/im. Conidia 
abundantly produced, arising singly, filiform, scolecosporous, arcuate to C- or S-shaped or 



*Santesson (1952:52) attributes combinations of these three epithets into Pyrenotrichum to Hohnel; Hohnel 
( 1 9 1 : 652) suggested these species should be placed in that genus but did not validly publish the transfers (they were 
also not accepted in his 'Index', lot: ciV. 678). 



LICHENICOLOUS COELOMYCETES 



61 



' *-£**. LW* '^J 




l.~* 




i -. 



*JL <Si~: " : 




r F *s 



Fig. 30 Pyrenotrichum splitgerberi (A-B, D-F, IMI 44258/; C, IMI 246129). A-B, Dark 
conidiomata (x 16). C, Light conMiomata (x25). D, Vertical section of conidoma showing 
portion with conidiogenous cells (left) and papillae not bearing conidia (centre to right) (x 1 020). 
E, Vertical section of conidioma (x 250). F, Vertical section of lower part of conidioma showing 
the branched conidiophores (x 1 020). 



62 



D. L. HAWKSWORTH 




Fig. 31 Pyrenotrichum splitgerberi (IMI 44258/). A, Conidiophores, conidiogenous cells 

and developing conidia. B, Conidia. 



otherwise convoluted, slightly thicker at the rounded apex, truncated and to about 1 jam 
wide at the extreme base, hyaline, multiseptate, with to 8 septa when mature, 
often with numerous guttules, smooth-walled, 55-80 x 2-3 jum. 

Hosts: According to Santesson (1952:40) this species is restricted to the thalli of 
foliicolous lichens belong to the genera Lopadium Korber (9 species) and Tapellaria Mull. 
Arg. (6 species). Parasymbiotic, sometimes apparently depressing the formation of apothecia 



LICHENICOLOUS COELOMYCETES 63 

in the host and then appearing as if a lichen-forming coelomycete. This species has been 
considered to be a lichen by several authors (e.g. Mameli-Calvino, 1930; Rizzini, 1952) and 
it may be that in some cases the fungus can take over the algae from the original lichen 
thallus it invades, so becoming lichenized (cf. Blarneya hibernica D. Hawksw. et al.\ 
Hawksworth et al, 1980). A critical anatomical investigation of the fungus-alga relationship 
in material of this species on 'sterile' thalli lacking apothecia would be of interest. 

Distribution: Probably pantropical. There are reliable reports at least from Brazil, China, 
Cuba, Ghana, New Zealand, Surinam, and the U.S.A. (Louisiana). 

Observations: I have not carried out a critical investigation into the range of variation of 
this species in view of Santesson's unpublished studies; the above description is mainly based 
on IMI 442587 (determined by Santesson in 1951). The synonymy presented above is derived 
from Keissler (1927, 1933), Santesson (1952 : 49-53), and a few minor sources; I have not 
studied any of the relevant type collections. Santesson (1952 : 41) mentions only one other 
species of the genus as having ± filiform unbranched and multiseptate conidia, P. filiferum 
R. Sant. ined., known from two species of Lopadium; the characters separating this species 
from P. splitgerberi were not indicated. 

Specimens: Brazil: Rio de Janeiro, Jardin Botanico, on Buxus sempivirens, 1947, C T. Rizzini 11 
(PC!); Parana, Desvio Ypiranga, 26 September 1909, P. Dusen 8822 (PC!); sine loc, comm. March 
1951, comm. R. Santesson (IMI 45098!).— Ghana: Pokoasi, on Diospyros sp., 1 June 1949, S. J. 
Hughes 1040 (IMI 43725c); loc. cit., on Lopadium sp. on Pancovia bijuga, 25 May 1949, S. J. Hughes 
834 (IMI 44258J!); Aburi, on Trycalysia cf. pallens, 24 May 1949, S. J. Hughes 807 (IMI 44170d!); 
near Nsuaem, 8 May 1949, S. J. Hughes 219 (IMI 44219b!); Togoland, Jasikan, on Cola caricifolia, 27 
May 1949, S. J. Hughes 957 (IMI 43662g!); Togoland, Hobae, on Culcasia scandens, 28 May 1949, 
S. J. Hughes 934b (IMI 44213a!).— U.S.A.: Louisiana, Assumption Parish, 2-3 mis W of Grand Bayou, 
on Sabal palmetto, 8 February 1976, S. C Tucker 1 5448 (IMI 246129!). 

2. Pyrenotrichum staurosporum (Keissler) R. Sant., Symb. bot. upsal 12(1): 41 (1952). 
(Fig. 32A-B) 

Chlorocyphella aeruginascens var. staurospora Keissler, Annln naturh. Mus. Wien 41 : 162 (1927). 

Icones: Keissler, Annln naturh. Mus. WienAX : 161 fig. lc (1927). 

This fungus is very similar in habit to Pyrenotrichum splitgerberi but is clearly separated 
from that species in that the conidia are branched with 2-4 arms arising at a single locus from 
a point about halfway along the total length of the conidium. In the specimens studied, the 
conidia were (overall) 60-85 x 2-5-3*5 jum. 

Hosts: According to Santesson (1952 : 40) this species is restricted to foliicolous lichens 
belong to the genus Lasioloma R. Sant. (4 species). Parasymbiotic. 

Distribution: This taxon was first described from Guam but I have seen specimens also 
from Ghana and Malaysia. 

Observations: This species is clearly separated from Pyrenotrichum splitgerberi on the 
basis of its branched conidia as well as the disparate hosts. Santesson (1952 : 4 1 ) transferred a 
second species with branched conidia to Pyrenotrichum, P. foliicola (Vainio) R. Sant.*; this 
was first described from the Phillipine Islands and occurs on foliicolous species ofBacidia de 
Not. (5 listed by Santesson, loc. cit.: 40). I have not seen any material of P. foliicola. Although 
Santesson did not give the basionyms of P. foliicola nor P. staurospoum, his combinations 
are treated as validly published here as these taxa were clearly accepted as distinct, albeit in a 
Table. As Santessons's monograph appeared before 1 January 1953, Art. 33.2 does not 
apply. 



*Symb. bot. upsal. 12(1): 41 (\952).—Cyphella foliicola Vainio, Annls Acad. Sci. fenn. 15(6): 83 (1921). 
Chlorocyphella foliicola (Vainio) Keissler, Annln naturh. Mus. Wien 41 : 1 59 (1927). 



64 



D. L. HAWKSWORTH 




Fig. 32 Pyrenotrichum staurosporum (IMI 444 \0d). A, Conidiogenous cells and developing 

conidia. B, Conidia. 

Specimens: Ghana: Aburi, on Landolphia owariensis, 24 May 1949, S. J. Hughes 791 (IMI 
444 \0d). — Malaysia: Cameron Highlands, on Lasioloma arachnoideum on Xanthophvllum affina, 6 
September 1953, W. J. Cherwick [A. Johnston no. 1113] (IMI 54915/-;?!). 



XIX. VOUAUXIELLA Petrak & H. Sydow 

Beik Repert. now Spec. Regni vegAl : 482 (1927). 
Alysia Cavalc. & Silva, Publcoes Inst. Micol. Recife 647 : 32 (1972). 



LICHENICOLOUS COELOMYCETES 65 

Conidiomata pycnidial, arising singly, subglobose to almost cupuliform, immersed or 
semi-immersed, dark brown to black; walls composed of thick-walled dark brown to 
olivaceous green cells forming a textura angularis but sometimes almost hyphal below, the 
upper cells separating to form an irregularly delimited ostiole. Conidiophores short, 
branched only at the base, septate, pale brown, smooth or verruculose, lining the inner wall 
of the pycnidium. Conidiogenous cells integrated or discrete, terminal, holothallic, 
subhyaline to pale brown or olivaceous green, smooth or verruculose, a basal thicker-walled 
'mother' cell present in some species and becoming flared at the onset of conidiogenesis. 
Conidia catenate, forming in acropetal chains, chains simple, with up to 8 or more adhering 
conidia, ellipsoid with both ends abruptly truncated (except in the apical conidium and in V. 
uniseptata), usually simple, rarely 1 -septate, olivaceous green, brown or dark brown, 
smooth or verruculose, liberated by the breakup of the chains, dry. 

Type species: Vouauxiella verrucosa (Vouaux) Petrak & H. Sydow (lectotype; selected by 
Clements & Shear, 1931 : 378). 

Number of species: Four, all obligately lichenicolous, and one of which may eventually 
prove not to be congeneric. 

Observations: Vouauxiella has an especially interesting method of conidiogenesis 
(Morgan-Jones, 1971; Sutton, 1980); indeed it is the only non-stromatic coelomycete with 
holothallic conidiogenesis recognized, with the possible exception of the new genus 
Nigropuncta described above (p. 46). As noted by Hawksworth (1978a : 185) and Sutton 
(1980 : 24), Vouauxiella uniseptata may well merit recognition as a distinct genus on the 
basis of the absence of conidiophore mother cells, much more deeply pigmented and 
1 -septate conidia, and the broadly opening pycnidia. Vouauxiella does, however, provide a 
convenient temporary niche for that fungus pending more detailed studies on conidiogenesis. 

Key to the species 

1 Conidia simple 

Conidia 1-septate, (10— )12— 1 5(— 17) x 5-7(-8)/^m, dark brown, smooth to weakly 

verruculose; pycnidia almost cupuliform, 40-70 //m diam . Vouauxiella uniseptata (p. 66) 

2 Conidia smooth-walled 

Conidia distinctly verruculose, 6-9 x 3*5—5 jum .... Vouauxiella verrucosa (p. 67) 

3 Conidia (5-)6-8(-9) x 3-4 //m; pycnidia mainly 60-90 //m diam 

Vouauxiella lichenicola (p. 65) 
Conidia 3—3-5 x 2'5//m [but see below]; pycnidia 70-160//m diam 

Vouauxiella pithospora (p. 66) 

1. Vouauxiella lichenicola (Lindsay) Petrak & H. Sydow, Beih. Repert. nov. Spec. Regni veg. 
42:484(1927). 

This species was described and illustrated in colour in great detail by Lindsay (1869a : pi. 
23 figs. 1-18), although he also included some other taxa in his concept. Its conidiogenesis is 
treated by Morgan-Jones (1971), and illustrations have been published more recently by 
Hawksworth (1976 : 58) and Sutton (1980 : 25). This well-known fungus is not therefore 
described in detail here because its diagnostic features will be apparent from the generic 
description and key. 

The typification and synonymy of Lindsay's epithet is discussed by Hawksworth 
1979:287). 

Hosts: Lecanora species, apothecia. Especially frequent on L. chlarotera Nyl. but reported 
from a range of other red-fruited species of the genus. The pycnidia of this species and 
Vouauxiella verrucosa are characteristically found at the edges of the disc adjoining the 
thalline margins; infected discs are commonly paler in colour than is usual for the lichen and 
piebald in appearance. Reports from other host genera are dubious, partly due to confusion 



66 D. L. HAWKSWORTH 

with Lichenodiplis lecanorae, a very different fungus (p. 38), to which taxon references to it 
in Keissler (1930 : 568) on at least Calopaca cerina (Ehrh. ex Hedw.) Th. Fr., Lecanora 
dispersa (Pers.) Sommerf., and Pertusaria leioplaca DC, should be referred. The report of V. 
lichenicola on Caloplaca holocarpa (Hoffm.) Wade by Werner (1969 : 204) may also belong 
to that fungus. 

Distribution: Probably widespread, at least in Europe. Reliably reported from the British 
Isles (England, Ireland, Scotland), France (Vouaux, 1914:297), Germany, Morocco 
(Werner, 1970, 1972), Spain (Santesson, 1960 : 519), Sweden (Santesson, 1949 : 143) and 
the U.S.A. (Mass.; Santesson, 1960 : 519). Material from Finland distributed under the name 
V. lichenicola by Rasanen (Lich. Fenn. Exs. no. 1 50, BM!) is a Taeniolella species. 

Specimens (all on Lecanora chlarotera apothecia): British Isles: S. Devon, Slapton, Slapton Ley 
Nature Reserve, 30 July 1973, D. L. Hawksworth 3395 (IMI 178325!); he. cit, 10 October 1974, D. L. 
Hawksworth 3930 (IMI 188759!); he. cit., Peasdish, 23 August 1980, D. L. Hawksworth 5044c (IMI 
251263!); Oxfordshire, Hook Norton, January 1973, H. J. M. Bowen (IMI 185887!); Sussex, Eridge Old 
Park, 20 May 1972, F. Rose (E!). Scotland, Kincardineshire, The Burn, Fettercairn, 26 July 1972, B. J. 
Coppins et al. (E!); Perthshire, Loch Tay, June 1856, W. L. Lindsay (E — lectotype!); E. Lothian, near 
Dunglass, Cockburnpath, June 1 856, J. M. Lindsay (E!). 

2. Vouauxiella pithospora (Cavalc. & Silva) B. Sutton, The Coelomycetes: 24 (1980). 
Alysia pithospora Cavalc. & Silva, in Cavalcante et al, Publgoes Inst. Micol. Recife 641 : 34 (1972). 

For description and illustrations see Cavalcante et al. (1972 : 36 fig. 7). 

Host: Originally described as a lichenized coelomycete with a Phycopeltis phycobiont, but 
almost certainly a lichenicolous species on an unknown foliicolous lichen. 

Distribution: Brazil (Rondonia). Known only from the original locality. 

Observations: Neither Sutton (loc. cit.) nor myself have seen material of this taxon but the 
published figures leave little doubt that a species of Vouauxiella was involved. The pyenidia 
and conidia were originally given as 70-160 jum diam and 2-6 x L5-2'5 jum, respectively. 
These measurements are not in accord with the scale on the figures which suggest the 
pyenidia are to about 60 pm wide and the conidia 3-3*5 x 2*5 pm; these latter figures were 
used by Sutton. It seems impossible to resolve this discrepancy in the absence of the type 
material. If the scale on the figure is correct, the species is clearly separated from the others 
treated here by the small conidia; if the text is correct it will merit a critical comparison with 
V. lichenicola. 

3. Vouauxiella uniseptata D. Hawksw., Notes R. bot. Gdn Edinb. 36 : 195 (1978). 

For description and illustrations see Hawksworth (1978a : 195—196), and Sutton 
(1980:24-25). 

Host: Parmelia laevigata (Sm.) Ach., thallus. Forming distinctive raised pustular black 
patches, mainly 1-2 mm diam, on the lobes. 

Distribution: British Isles (Scotland). Three collections were reported by Hawksworth {loc. 
cit.) and four others have since been seen (see below). Not yet seen by me on this host in 
Devon despite careful searches. 

Observations: This species may well eventually require transfer to an independent genus 
for the reasons indicated above (p. 65). 

Specimens (additional to those listed by Hawksworth, loc. cit.): British Isles: Scotland, Argyll, S side 
Loch Sunart, 14 August 1972, S. R. Davey (IMI 250032a!), Benderloch, grounds of Lochnell House, 4 
August 1980, B. J. Coppins 8052 (E, IMI 251027!), N side Loch Etive, 8 km E of Connel Burn, 7 
August 1980, B. J. Coppins 8083 (E, IMI 251025!); Kirkudbright, Kintyre, Tarbert, Wilhashemraig, on 
Alnus, 29 July 1973,7. E. Menlove (IMI 228667!). 



LICHENICOLOUS COELOMYCETES 67 

4. Vouauxiella verrucosa (Vouaux) Petrak & H. Sydow, Beih. Repert. nov. Spec. Regni veg 
42:483(1927). 

For illustrations of this species and discussion of its separation from V. lichenicola see 
Hawksworth(1976 : 58). Further illustrations are included in Sutton (1980 : 25). 

Host: Lecanora chlarotera Nyl. and L. laevis Poelt, apothecia and rarely thallus. The 
pycnidia have a similar habit to those of Vouauxiella lichenicola as described above and 
microscopic examination is therefore needed to separate these two species. 

Distribution: British Isles (England, Ireland, Scotland), France and Pakistan. Perhaps often 
overlooked as V. lichenicola and perhaps much under-recorded, at least in Europe. 

Observations: Lindsay (1869a) included material of this taxon within his concept of V. 
lichenicola but, in accordance with later usage, a smooth-spored specimen was selected as 
lectotype for his epithet (see above). 

Specimens (additional to those listed by Hawksworth, be. cit.): British Isles: England, Wiltshire, 
Imber, on Lecanora chlarotera, 1979, A. M. Burnet (IMI 237278!). Ireland, Co. Cork, near Cork, on L. 
chlarotera, March 1858, /. Carroll (E!). — Pakistan: Khanaspur, on L. laevis, 25 April 1975, S. Ahmad 
1059 (IMI 199857c!). 

XX. VOUAUXIOMYCES Dyko & D. Hawksw. 

in Hawksworth & Dyko, Lichenologist 11 : 57 (1979). 

For a description and discussion of the affinities of this genus see Hawksworth & Dyko 
(1979). 

Type species: Vouauxiomyces truncatus (B. de Lesd.) Dyko & D. Hawksw. 

Teleomorph (perfect state): Abrothallus de Not. In the course of investigations into the 
anamorphs of lichenicolous ascomycetes it became clear that those of Abrothallus were 
referrable to Vouauxiomyces. All connections are based on strong circumstantial evidence, 
the concurrence of ascomata and conidiomata, and have not been proved by ascospore 
cultures. 

Number of species: Originally described as monotypic, but two additional species are 
added here. These three anamorphs all regularly occur without the teleomorph and it is 
consequently useful to have binomials available for these phases. The genus Abrothallus has 
been studied intensively by Santesson but his results still await publication; according to 
Santesson (in Nordin, 1964 : 226) the genus comprises about 15 species and it is probable 
that others have anamorphs in Vouauxiomyces also. Published accounts of anamorphs in 
four species of Abrothallus in addition to the three treated here are noted on p. 72. 

Key to the species 

1 Conidia exceeding 6 //m in length 2 

Conidia 3-5(-6) x 2-3'5(^) //m; on Ramalina species . . Vouauxiomyces ramalinae (p. 67) 

2(1) Conidia 6- 5-8 x 4-5 -5 //m; on Parmelia caperata and P. perlata 

Vouauxiomyces truncatus (p. 70) 
- Conidia (7— )7"5— 10'5(— 1 F5) x (5-)5-5-7(-7*5)//m; on Parmelia saxatilis and Platismatia 

glauca Vouauxiomyces santessonii (p. 69) 

1. Vouauxiomyces ramalinae (Nordin) D. Hawksw. comb. nov. 
(Fig. 33A-B) 

Phoma ramalinae Nordin, Svensk bot. Tidskr. 58 : 227 (1964). 

Type: Sweden, Uppland, Malsta par., on Ramalina fraxinea (L.) Ach. (apothecia), 23 May 1 96 1 , /. 
Nordin 563a (UPS— holotype n.v.). 

Icones: Nordin, Svensk bot. Tidskr. 58 : pi. I figs 1-4, pi. II figs. 5 and 7 (1964). 



68 




D. L. HAWKSWORTI 

B 



OOOOOo 
0G0DQ0 





lOjLLm 

Fig. 33A-B, Vouauxiomyces ramalinae (IMI 194098); A, conidiogenous cells; B, conidia. 
C-D, V. santessonii (UPS — holotype); C, conidiogenous cells; D, conidia. E, V. truncatus 
(Savoie, Santesson, UPS), conidia. 



Conidiomata pycnidial, immersed at first but becoming erumpent, to about half exposed 
at maturity, arising singly, scattered or sometimes in a ring in young infections, black, 
ostiolate, 55—75 //m diam; pycnidial wall mainly 3-5 cells thick, thickest in the exposed 
part adjacent to the ostiole, 5-12 /im thick, dark brown, pseudoparenchymatous, textura 
angularis, cells thick-walled, tending to be polyhedral, mainly 3-5 /um diam. 
Conidiogenous cells holoblastic, ampulliform to lageniform, lining the pycnidial cavity, 



LICHENICOLOUS COELOMYCETES 69 

percurrently proliferating, to 3 annellations seen, hyaline, smooth-walled (4-)6-10x 
3-5 jum. Conidia arising singly, obpyriform, hyaline, collecting in a mucilaginous mass in 
the pycnidial cavity and extruded as a drop, simple, irregularly guttulate, apex rounded, the 
base abruptly truncate, ± smooth-walled by light microscopy but indications of a sparse 
perhaps echinulate ornamation are sometimes visible by interference contrast, 3-5(-6)x 
2-3-5(-4)//m. 

Teleomorph (perfect state): Abrothallus suecicus (Kirschst.) Nordin. The connection is 
based on the repeated occurrence of the ascomata in assocation with the anamorph. 

Hosts: Ramalina calicaris (L.) Fr., R. dilacerata (Hoffm.) Hoffm., R.fastigiata (Pers.) Ach. 
and R. fraxinea (L.) Ach., thallus and especially apothecia. These hosts are taken from 
Nordin (1964 : 228-229) who also indicated that the species might well be present on R. 
sinensis Jatta. The symptoms and course of infection were studied carefully by Nordin, who 
found the pycnidia to start at one point near the margin of an apothecial disc and then extend 
over the disc forming pycnidia concentrically, the discs become greyish and the surface 
layers extremely brittle. 

Distribution: British Isles (England, Ireland, Scotland), France, Finland, Norway and 
Sweden. Nordin (1964) reported Abrothallus suecicus from 104 localities, 50 of these as the 
anamorph alone. Evidently widespread where the host lichens are well-developed. 

Observations: Vouauxiomyces ramalinae is distinguished from the other species of the 
genus accepted here by the smaller pycnidia, the shorter and narrower conidia, and in being 
associated with a different teleomorph. As the pycnidia are not uncommonly found without 
the ascomata, it is clearly desirable that the anamorph should have its own binomial; 
Nordin's epithet is consequently recombined into Vouauxiomyces here. The anamorph 
cannot be accommodated in Phoma, as currently circumscribed, because the conidiogenous 
cells are holoblastic and annellate and the conidia are not regularly bacilliform, ellipsoid or 
subglobose. 

Specimens (see Nordin, 1964 : 229-230 for numerous other specimens he studied): British Isles: 
England, S. Devon, Slapton, Duck Marsh, on Ramalina fastigiata on Salix, 21 May 1975, D. L. 
Hawksworth 4137 (IMI 194098!). Ireland, Co. Galway, Connemara, Kilkieran, Flannery Bridge, on R. 
fastigiata, 1 August 1966, G. Morgan- J ones (IMI 1214426!). 

2. Vouauxiomyces santessonii D. Hawksw. sp. nov. 
(Fig. 33C-D) 

Similis Vouauxiomvcei truncato (B. de Lesd.) Dyko & D. Hawksw. sed difTert in conidiis latioribus, 
(7-)7-5-10-5(-ll-5)x'(5-)5-5-7(-7-5)//m. 

Typus: Suecica, Uppland, Alsike s:n, Lunsen, vid akertegen SV om Hjortronmossen, pa grenar pa 
marken i granskog, in thallo Platismatiae glaucae (L.) Culb. & C. Culb., 23. ix. 1947, R. Santesson 
(UPS— holotypus!). 

Icones: Kotte, Zentralbl. Bakt. ParasitKde II, 24 : pi. 3 fig. 32 (1909).— Lindsay, Q. Jl microsc. Sci. 
II, 7 : pi. 5 figs. 5-7 (1857).— Schaechtelin & Werner, Bull, trimest. Soc. mvcol. Fr. 42 : pi. 3 figs. 
11-12(1926). 

Conidiomata pycnidial, immersed at first but becoming erumpent through the surface of 
the host, about \ exposed at maturity, arising singly, scattered, sometimes arising on the site 
of a previous pycnidium and then assuming a rimmed appearance from above and a 
double-walled structure in vertical section, black, ostiolate, the ostiole often conspicuous 
and c. 25 jum diam, 125-225 jum diam; wall of varying numbers of cell layers, mainly 4-8, 
15-25//m thick, thickest around the ostiole in the erumpent part of the pycnidium, dark 
brown, pseudoparenchymatous, textura angularis, cells thick-walled, tending to be poly- 
hedral, mainly 3^//m diam. Conidiogenous cells holoblastic, ampulliform to lageniform, 
lining the pycnidial cavity, percurrently proliferating, annellate with to 4 annellations seen, 
hyaline, smooth-walled to weakly echinulate, 8-12 x 3-4 jum. Conidia obpyriform, hyaline, 



70 D. L. HAWKSWORTH 

collecting in a mucilaginous mass in the pycnidial cavity and extruded as a drop, simple, 
sometimes 1-3 guttulate, apex rounded, the base abruptly truncated, thin-walled, ± smooth- 
walled at lower magnifications but with a widely spaced echinulate ornamentation just 
discernible by interference contrast, (7-)7*5-10*5(-l 1*5) x (5-)5'5-7(-7-5)/zm. 

Teleomorph (perfect state): Abrothallus parmeliarum (Sommerf.) Arnold. The conidia of 
this species were well-described in some of the earliest papers on lichenicolous fungi, notably 
those of Tulasne (1852 : 1 14*) and Lindsay (1857*), and the connection has been accepted 
by subsequent students of Abrothallus (e.g. Kotte, 1909 : 87; Schaechtelin & Werner, 
1926:238-239). 

Hosts: Parmelia saxatilis (L.) Ach. and Platismatia glauca (L.) Culb. & C. Culb., thalli. 
The teleomorph is reported from a large number of species of Parmelia, many of which must 
also support the anamorph. In early infections the pycnidia arise on almost healthy lobes, 
with a circle of brownish discoloured tissue around the pycnidia (rarely over 25 //m thick). 
In more advanced stages of infection the pycnidia occur on gall-like deformations of the host 
thalli typical of infections due to Abrothallus parmeliarum. 

Distribution: I have studied material of the anamorph from the British Isles and Sweden, 
but it can be expected to occur throughout the range of the teleomorph. The distribution of 
the teleomorph is difficult to ascertain from the literature due to taxonomic uncertainties but 
Abrothallus parmeliarum s. lat., in addition to being widespread in Europe, is mentioned 
from at least Canada, Chile, Bolivia, Ecuador, Hawaii, Tenerife, Kenya, Tasmania, 
Morocco, New Zealand, South Africa, Mongolia, the U.S.A., Siberia and Venezuela (Hertel, 
1971 : 228-229). 

Observations: This species differs from Vouauxiomyces truncatus in the broader, and 
often also longer, conidia; the taxonomic importance of this difference is substantiated by the 
different teleomorphs and host ranges. 

The species is named in honour of Prof. Dr R. Santesson in recognition of his important, 
yet unpublished, studies on Abrothallus, Pyrenotrichum, and other lichenicolous fungi. 

The name Epicoccum parmeliarum H. Olivier may provide an earlier epithet for this 
species but is not taken up here in view of some uncertainties surrounding its application (see 
p. 76). 

Additional specimens: British Isles: Scotland, Argyll, 4 km E of Dalmally, wood on N side of River 
Lochy, on Parmelia saxatilis on Quercus, 1 August 1980, P. W. James & B. J. Coppins 4961 (E, IMI 
251031!). Wales, Montgomery, Gregynog Hall, on P. saxatilis on Fagus sylvatica, 25 May 1979, K. L. 
Brady (IMI 239067!). — Sweden (all on Platismatia glauca): Uppland, Vange s:n, Fiby urskog (S om 
Fibysjon), i granskogen 6 om Getryggen, pa grankvistar, 28 April 1948, R. Santesson 4859 (UPS!); 
Viksta s:n, Djupdalen, c. \ km V om Nyboda, pa gran i tat barrskog, 21 May 1947, R. Santesson 
(UPS!); Rasbo parish, c. 1 km SSE of the lake Rokarn, on twigs oiPicea abies in a rather dark forest, 23 
October 1955, R. Santesson 11139b (UPS!); Skuttunge, Norra myren, 3 May 1939, R. Sernander 
(UPS!); Uppsala, Vange socken, Fiby urskog, 15 November 1945, B. H. Svenonius (UPS!). 
Vastergotland, Halleberg, Vanersnas s:n, Hallesnipan, pa ek, 15 June 1944, R. Santesson (UPS!). 

3. Vouauxiomyces truncatus (B. de Lesd.) Dyko & D. Hawksw., in Hawksworth & Dyko, 
Lichenologist 11 : 58 (1979). 
(Fig.33E,34) 

For description, illustrations and synonymy see Hawksworth & Dyko ( 1979). 

Teleomorph (perfect state): Abrothallus microspermus Tul. The anamorph of this species 
was illustrated and described by Tulasne (1852:116, pi. 16 figs. 22-26) in his 
original account of the fungus, giving the conidia as 6*5 x3'5/im. Hepp (Flecht. Eur. no. 

*These reports were given under the name Abrothallus smithii Tul., a superfluous name for A. bertianus de Not. 
(which has smaller conidia; see p. 72) misapplied by these authors. 



LICHENICOLOUS COELOMYCETES 



71 




Fig. 34 Vouauxiomyces truncatus (Savoie, Santesson, UPS), conidia (x 2500). 

471, 1857; K!) later distributed and illustrated material of both states. Pycnidia commonly 
occur without the teleomorph but are regularly encountered mixed amongst the ascomata. 
Abrothallus microspermus differs from A. parmeliarum, with which it has often been 
confused, in the consistently epruinose ascomata which only reach 300 jum diam; according 
to Keissler ( 1930) the ascospores are also paler brown and the epithecium K- 

Hosts: Parmelia caperata (L.) Ach., and more rarely P. perlata (Huds.) Ach., thalli. 
Little damage is usually encountered and the fungus is probably parasymbiotic. 

Distribution: Austria, the British Isles (England, Ireland, Scotland, Wales), France, 
Germany and Switzerland. The report of the occurrence of this species in Sweden 
(Santesson, 1949 : 142) is based on material now referred to Vouauxiomyces santessonii. 

Specimens (additional to those listed by Hawksworth & Dyko, 1979): British Isles: England, 
S. Devon, Slapton Ley Nature Reserve, The Causeway, on Parmelia perlata on Salix, 30 March 
1972, D. L. Hawksworth 2696 p.p. (IMI 164972 p.p.l, UPS!); he. cit, Peasdish, on P. caperata 
on Ulmus, 23 August 1980, D. L. Hawksworth 5043 (IMI 251260!); Dorset, Wareham, on P. 
caperata, 1978, V. Giavarini (E!). Scotland, Dunbarton, Loch Lomond, Aber Island, on P. 
perlata, 10 October 1979, B. J. Coppins 4482 (E, IMI 243340!). Wales, Merioneth, Barmouth, 
nr Arthog, Fegla Fawr, on P. caperata, 12 December 1972, B. J. Coppins (E!). — France: 
Savoie, Dept. Haute-Savoie, N of Sciez, Chateau Coudree, on P. caperata on Tilia, 9 June 
1946, R. Santesson (UPS!).— Switzerland: sine loc, P. Hepp [Flecht. Eur. no. 471] (K!). 



Additional reports of anamorphs 

In addition to the anamorphs of Abrothallus and Microcalicium species treated in the 
preceding section, which can occur without their teleomorphs, there are literature 
reports of pyenidial anamorphs in a number of other lichenicolous ascomycetes. As 
many of these may be merely spermatial, the conidia not functioning as diaspores but 
rather as gametes, and consequently are normally to be found in association with 



72 D. L. HAWKSWORTH 

ascomata, it seems superfluous to provide them with independent binomials. No 
compilation of the lichenicolous ascomycetes in which such anamorphs have been 
reported appears to be available. As these states may cause confusion when studying 
lichenicolous Coelomycetes, or sometimes be found alone, reports of those I have 
been able to discover are listed here. The location of all such reports would be a major 
bibliographical exercise involving the study of the whole literature on lichenicolous 
fungi; the following should consequently not be viewed as an exhaustive list. 

Abrothallus bertianus de Not. (syn. A. glabratulae Kotte, A. smithii Tul.) 

Kotte (1909:81) gave the conidia in this species as 5'46-6*5 x 39-^'42 /im, but I 
have not found a pycnidial state in the few specimens of it I have seen. This fungus 
appears to be confined to brown species of Parmelia and is especially frequent on P. 
glabratula (Lamy) Nyl. (Santesson, 1960; Hawksworth & Minter, 1980). 

Abrothallus cetrariae Kotte 

In the original account of this species, described from Platismatia glauca, the 
conidia were given as 4'94-6 - 5 x 3'9-4'9 //m (Kotte, 1909:84). These measurements 
are too small for Vouauxiomyces santessonii (p. 69), which occurs on the same host. 

Abrothallus peyritischii (B. Stein) Kotte 

The pycnidia of this species, which occurs on Cetraria nivalis (L.) Ach., were 
illustrated by Kotte (1909: pi. 1 figs. 12-17) and appear to belong to Vouauxiomyces', 
the conidia were said to measure 5'2-6'76 x 39— 5*2 /zm. One of Kotte's figures is 
reproduced by Keissler(1930 : 205 fig. 46). 

Abrothallus usneae Rabenh. 

This fungus, which has often been united with A. parmeliarum, was investigated 
anatomically by Galloe (1950:73, pis. 113-115). His drawings of vertical sections of 
pycnidia, conidiogenous cells and conidia leave no doubt that the pycnidia of this 
species belong to Vouauxiomyces, but no conidial dimensions were provided. 

Arthonia clemens (Tul.) Th. Fr. (syn. Conidia clemens (Tul.) Massal., Phacopsis clemens 

Tul.) 

In the original description of this species, based on material from species of 
Lecanora and Rhizoplaca chrysoleuca, Tulasne (1852:125) described the pycnidia 
as 70-80 /im tall, thick- and brown-walled, with straight, bacilliform conidia 3-4 jum 
long. They were also reported from this fungus in the hymenium of Lecanora atra 
(Huds.) Ach. by Werner &Deschatres( 1974 : 317-8) who gave them as 4 x05 jum. 

Arthonia fuscopurpurea (Tul.) R. Sant. (syn. Celidium fuscopurpureum Tul.) 

This species, which occurs on Peltigera thalli, is described as forming bacilliform 
conidia which are hyaline and to 3*5 p.m long (Tulasne, 1852 : 121). 

Arthonia varia (Tul.) Jatta (syn. Phacopsis varia Tul.). 

Tulasne (1852:126, pi. 14 fig. 3) described straight bacilliform conidia about 
4'5 fim long in this species on Xanthoria parietina (L.) Th. Fr. The relationship of 
this taxon to both A. epiphyscia Tul. and A. destruens Rehm, which also attack 
Xanthoria species, requires further study; contrary to Keissler's (1930 : 98) treatment, it is 
distinct from A. glaucomaria (Nyl.) Nyl. which is found on Lecanora rupicola (L.) Zahlbr. 

'Celidium' insidens Vouaux 

The pycnidia of this fungus were rather fully described by Vouaux (1914:170). 
They arose in groups of 5-6 and were lenticular-convex, immersed only at the base, 
02-0-3 mm diam, black, and produced cuneiform hyaline 1 -septate conidia 9-12 x 4*5 jum 



LICHENICOLOUS COELOMYCETES 73 

from simple to 2-septate conidiogenous cells measuring 9-20 x 2-3 pim. The species was 
found on material of Ramalina fraxinea collected in Tenerife. 

Dacampiosphaeria rivana (de Not.) D. Hawksw. (syn. Homostegia lichenum Fuckel) 

According to Fuckel (1870:224) this fungus produces 1 -septate, cylindrical, curved 
conidia which are 16x2*5//m. In my investigations on this fungus I did not 
find any pycnidial state (Hawksworth, 1980a : 367—371), and it seems probable that 
Fuckel's report is based on an unidentified lichenicolous coelomycete on Peltigera. 

Dichosporium glomerata Pat. 

This species, which occurred on an undetermined corticolous lichen from Guadaloupe, is 
described as producing pycnidia on a superficial mycelium which give rise to hyaline, 
simple, subcylindrical conidia, measuring 5-6 x 2 jum (Vouaux, 1912 : 199). 

Didymella parvispora Henssen 

Henssen (1977:44, pi. 8k), in the original description of this fungus from 
Zahlbrucknerella africana Henssen, reported the pycnidia as partially immersed, 
elongate, 30-50 //m broad, with a dark brown ostiole, and elongate conidiogenous cells 
giving rise to bacilliform, hyaline conidia 1 *5— 2 x 0*5—1 //m. 

Guignardia olivieri (Vouaux) Sacc. 

Pycnidia resembling those of Leptodothiorella, scattered in thalline warts, immersed, ± 
globose, 75-100 jum diam; pycnidial wall subhyaline, pseudoparenchymatous, 8-12 jum 
thick; conidiogenous cells phialidic, short-ampulliform, 7—10 x 1*5-2*5 //m; conidia arising 
in short chains, bacilliform, hyaline, simple, often slightly compressed medianly, 3*5-5 x 
1*5-2 jum. The anamorph of this species was described and illustrated by Hawksworth 
(1975/?: 190-191, fig, 5g-j) on Xanthoria parietina. The conidiogenous cells were then 
described as holoblastic, but improved microscopic techniques indicate that they are almost 
certainly enteroblastic-phialidic. 

Homostegia piggotii (Berk. & Broome) P. Karsten 

This species was studied in some detail by Schaechtelin & Werner (1928) who 
noted pycnidia in developing stromata on Parmelia saxatilis (L.) Ach. These were 
pyriform, ostiolate, with slender conidiogenous cells radially orientated. Their illustration 
(loc. cit.: pi. 1 1 fig. 17) suggests that the conidiogenous cells may sometimes be multicellular 
and branched, but the hyaline, simple, subglobose conidia were reported to be formed 
apically. No measurements were provided. 

Lethariicola sipei Grumm. 

In the original account of this monotypic genus of odontotremoid fungi on Letharia 
vulpina (L.) Hue, Grummann (1969: pi. XIV figs. 19-20) provided photographs of 
rather thick-walled erumpent pycnidia, but did not mention them in the written 
account of the species. The nature of the conidiogenous cells and the conidia cannot 
be deduced from the low-power photographs provided. 

Lichenomyces lichenum (Sommerf. ex Fr.) R. Sant. (syn. Celidium stictarum Tul.) 

Pycnidia have been noted in this species by several workers, notably Tulasne 
(1852: 123, pi. 14 fig. 8), Werner (1928: pi. IX figs. 2-4) and Vezda (1970:222, 225 
fig. 4). The pycnidia are immersed to slightly erumpent at the ostiole in young 
stromata, and contain filiform, simple conidiogenous cells to 30 //m tall, which in 
turn produce bacilliform, hyaline, simple conidia 3^ x 1 jum singly from their apices. This 
fungus is especially frequent on Lobaria pulmonaria (L.) Hoffm., and is known from a wide 
range of genera in the Stictaceae. 



74 D. L. HAWKSWORTH 

Merismatium lecanorae (H. Olivier) Vouaux 

Vouaux (1913:78) discovered material of this fungus, on Ochrolechia parella (L.) 
Massal., with pycnidia which were black, 250-350 jum diam, and formed ovoid 
conidia which were hyaline and simple at first, but later became brown and 3-septate 
to muriform when mature, and 1 1-15 x 6-7 jum. He was not, however, convinced of 
the relationship between the pycnidia and the putative teleomorph. 

Muellerella lichenicola (Sommerf. ex Fr.) D. Hawksw. (syn. Tichothecium erraticum 
Massal.) 
Vouaux (1913:63) found pycnidia in a collection of Caloplaca vitellinula auct. 
associated with the species now called Muellerella lichenicola. These gave rise to 
brown, 1 -septate, ellipsoid conidia measuring 6-7 x 2*5—3 //m. Vouaux had little doubt that 
he was dealing with an anamorph of M. lichenicola, but the details he provides show that his 
material belonged to Lichenodiplis lecanorae (see p. 38). 

Nesolechia oxyspora (Tul.) Massal. 

Kotte (1909:93, pi. 3 figs. 29-30) noted pycnidia in the thecium of this species, 
which normally occurs on thalli of Parmelia saxatilis. These produced rather irregularly 
shaped broadly ellipsoid to subglobose hyaline conidia, but no further details were provided. 
He may have been dealing with an unknown fungicolous coelomycete on the Nesolechia, 
rather than an anamorph of N. oxyspora. 

Scutula epi Waste matica (Wallr.) Rehm (syn. S. wallrothii Tul.) 

According to Tulasne (1852: 119-120, pi. 14 figs. 18-24) and Vouaux (1913:424), 
this fungus, which occurs on Peltigera thalli, has two types of pycnidia. One, which 
they considered to be spermagonial, had pycnidia 60-90 //m diam with hyphal walls 
and formed cylindrical, slightly curved, simple, hyaline conidia, 8-12 x 0*5-1 jum; 
the second type of pycnidium was larger, 100-1 50 jum diam, with the walls composed 
of thick-walled pseudoparenchymatous cells, and gave rise to narrowly ellipsoid, 
often slightly curved, simple or rarely 1-septate, hyaline conidia, 12-16 x 2- 5-^4 //m, 
from conidiophores measuring 6-10 x 2 jum. There is a strong resemblance between 
the reported macroconidial phase and the description of Diplodina peltigerae (see 
p. 76). The pycnidial wall structure for this assumed state, as illustrated by Tulasne 
(loc. cit.), recalls that of Karsteniomyces peltigerae (see p. 22) and it is conceivable that he 
had a mixed collection in which the conidia of the Karsteniomyces had not attained their 
normal length and most not their final septum. 

Stigmidium dispersum (Lahm ex Korber) D. Hawksw. (syn. Pharcidia dispersa (Lahm 
ex Korber) Winter) 
Pycnidia were apparently first reported for this species by Vezda (1963:151-153, 
fig. 4), who described them as semi-immersed, 80-120 jum diam, and forming bacilli- 
form, simple or 1-septate, hyaline conidia, 10-15 x 15 //m. 

Telimena' foreaui F. Moreau 

In the original account of this species, described from specimens of Heterodermia 
speciosa (Wulfen) Trevisan collected in India, Moreau (1951:209) reported that 
pycnidia were present in the stroma. These were c. 50 //m diam, immersed, and 
formed bacilliform, hyaline conidia, 4 x 1 jum. 



Excluded species 

All taxa referred to coelomycete genera which have been considered as lichenicolous 
by their original or later authors, and which have not been treated in the preceding 



LICHENICOLOUS COELOMYCETES 75 

sections of this contribution, are compiled here alphabetically; obligate synonyms 
are listed by their basionyms and not separately. 

Aposphaeria ramalinae Vouaux, in Pitard & Harmand, Bull. Soc. bot. Fr., Mem. 
22 : 72(1912)[ fc 191 1']. 

Type: Canary Islands, Tenerife, Vueltas de Taganana, *sur le Ramalina implectens, dans 
la zone sylvestre 1 , alt. 700 m, 1907, J. Pitard. 

This fungus was originally described as forming a brownish superficial reticulum of 
hyphae 1*5-3 jam wide, and from which elongate fusiform brownish l(-3) septate 
conidia 9-12 x 2-3*5 jam were formed in short chains. In the centre of the mycelium 
pycnidia 80-1 30 jum were produced which contained hyaline, simple, broadly ovate 
conidia, measuring 3-3*5 x 2*5 jum. No material of this taxon could be located amongst 
Harmand's herbarium in Angers (M. Guerlesquin, in litt), but according to the original 
description the name seems to have been based on two different fungi: an unidentifiable 
dematiaceous hyphomycete and a coelomycete. The conidia of the latter recall those of 
Vouauxiomyces ramalinae (see p. 67), but the pycnidia were larger than usual for that 
species. 

Chaetothyriolum puiggarii Speg., Boln Acad. nac. Cienc. Cordoba 23 : 522 (1919). 

This monotypic genus was considered by Petrak & Sydow (1935 : 192) to be based 
on the pycnidia of a lichen. The type material was re-examined by Santesson 
(1952 : 53) and found to be too fragmentary for reliable determination. 

Chlorocyphella aeruginascens var. cystidiifera Keissler, Annln naturh. Mus. Wien 41 : 162 
(1952). 
This fungus, described from material of Lopadium phyllogenum (Mull. Arg.) Zahlbr. 
collected in the Solomon Islands, is almost certainly a species of Pyrenotrichum. According 
to Keissler (loc. cit.), it differs from the species now known as P. split gerberi in having sterile 
cystidia-like structures between the conidiogenous cells, and much longer conidia, 
120-140 x 15 //m. I have not examined the type material and the name was not discussed 
by Santesson (1952 : 41 , 49-53). 

Coniothyrium epiphyllum Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 296 (1914). 

This name is of uncertain application, no fungus recalling the original description 
being present on the holotype (Hawksworth, 1977 : 193). 

Coniothyrium harmandii Zopfex Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 295 (1914). 

This name is of uncertain application in the absence of type material (Hawksworth 
1977: 193-194). 

Coniscosolen mirabilis Schilling, Hedwigia 67 : 297 (1927). 

This monotypic genus was based on the normal pycnidia of an unidentified 
Psorotheciopsis Rehm species according to Santesson (1952 : 331) who examined the type 
material. 

Cristidium pallidum R. Sant., Symb. bot. upsal. 12(1) : 357 (1952); nom. inval. (Art. 32). 

This name was mentioned as an undescribed genus and species of imperfect fungi 
on Gyalectidium ftlicinum Mull. Arg., but no further details were given and it is thus 
uncertain if this was a coelomycete or hyphomycete. It is conceivable that this might 
be Hansfordiellopsis lichenicola (Bat. & Maia) Deighton, which is well known on 
Gyalectidium (Hawksworth, 1979 : 224-227). 

Dendrophoma lecanorae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 284 (1914). 

This species was described on the basis of two different French collections, neither 
present among the remnants of Vouaux's herbarium (Rondon, 1970); on Lecanora 
piniperda Korber (syn. L. glaucella (Flotow) Nyl.) from Meyrueis, Lozere, and on 



76 D. L. HAWKSWORTH 

Lecania fuscella (Schaerer) Korber (syn. L. syringea (Ach.) Th. Fr.) from Malo- 
Terminus. The erumpent pycnidia were described as 100-172 //m, with a hyphal 
wall, 2-3-irregularly branched conidiophores 12-25 x 2 //m, and ovoid hyaline simple 
conidia 3-5 x 1-5-2 //m. In the absence of material this name remains of uncertain appli- 
cation; the conidiophore description and conidium shape indicates that Lecania conidia 
were probably not involved (Eigler, 1969 : pi. 5). 

Diplodia parmeliae Berk. & M. A. Curtis, in Berkeley, Grevillea 3 : 3 (1874). 

Type: U.S.A., South Carolina, on Parmelia cf. rudecta Ach., M. A. Curtis (K — holotype!). 
Diplodina parmeliae (Berk. & M. A. Curtis) Sacc, Syll. Fung. 3 : 413 (1884). 

This species was introduced with an extremely brief description which included no 
information on the sizes of spores or other features. The perithecia were reported to 
have depressed ostioles and to be loosely clothed with flexuous threads, but their 
colour was not indicated, and the spores stated to be hyaline with 1 septum and 
unequal cells. Examination of the holotype established that the perithecia were translucent- 
orange, about 200 fim. wide, with whitish lax hairs, and that ascospores in two size ranges 
were formed, macroascospores about 51x23/zm and microascospores 9-llx4-6//m. 
The fungus agrees in all details with Nectria heterospora Speg., a species described and 
discussed by Hawksworth & Booth (1976), but as the epithet heterospora dates from 1889, 
the new combination Nectria parmeliae (Berk. & M. A. Curtis) D. Hawksw. comb. nov. must 
be adopted for this species. 

Diplodina peltigerae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 289 (1914). 

Type: France, Vosges, Plainfaing, on Peltigera canina (L.) Willd., J. Harmand & L. 
Vouaux. 
This species is not represented in the remnants of Vouaux's herbarium (Rondon, 
1970) and it could not be located amongst Harmand's collections in Angers (M. 
Guerlesquin, in litt.). It was described as forming black, immersed pycnidia 60-100 jum 
diam, which produced hyaline, ellipsoid conidia with a single septum often located below 
the centre and 10-14 x 3-4 jum. If it is re-collected it should be possible to recognize it from 
these details and then re-assess its position. No similar fungus is known to occur on Peltigera 
(Hawksworth, 1980a); the measurements are suggestive of the reported macroconidial 
anamorph ofScutula epiblastematica (p. 74). 

Epicoccum parmeliarum H. Olivier, Bull internat. geogr. Bot. 17 : 232 (1907). 
Type: France, Orne, on Parmelia caperata and P. sulcata (thalli), H. Olivier. 
Phoma parmeliarum (H. Olivier) Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 196 (1914). 

Vouaux (loc. cit.) considered that this name was based on the anamorph of Abrothallus 
parmeliarum. There can be little doubt Olivier was dealing with a Vouauxiomyces species 
since the conidia were originally described by him as 'terminees en pedoncule a la base'; the 
pycnidial size quoted (0-2-0*4 mm diam) and the lengths and breadths of the conidia 
(6-9 x 5-7 jum) certainly recall the measurements for V. santessonii (teleomorph A. 
parmeliarum), but the mention of Parmelia caperata suggests that he may really have been 
dealing with either V. truncatus (teleomorph A. micro spermus), or a mixture of both these 
anamorphs on different hosts. As Olivier's epithet would pre-date both " santessoniP and 
'truncatus' at species level it seems unwise to take it up here in either sense in the absence of 
Olivier's material. 

Libertiella obscurior Gerber, Arch. Protistenk. 74 : 485 (193 1 ). 

Type: Austria, Hohen Tauren [Hohentauren], Naturschutzpark, on Hypogymnia intestiniformis 
(Vill.) Rasanen (syn. H. encausta (Sm.) W. Watson), 1929, F. Mattick. 

The holotype was formerly preserved in the herbarium at DR but the lichen 
material there was destroyed in 1945 and no duplicate appears to have existed (F. 



LICHENICOLOUS COELOMYCETES 



*A 






A 






v >- 






# 






77 











Fig. 35 Lichenophoma haematommatis. A, Pycnidia on the host. B, 
pycnidium. Scale not indicated. Reproduced from Keissler (1911 : 297). 



< Q->f '■'*■> - '•* r' ■■'•J* " 

Vertical section of 



Mattick, in litt.). This taxon was described as producing numerous immersed pycnidia, 
0* 1 5-0*2 mm diam, pale at the base but brownish above, delicate and hyphal, and producing 
ellipsoid, hyaline conidia measuring 2*5-3*4 x 1*7-2 jiva. No conidiophores were recog- 
nized. Pycnidia are often numerous in Hypogymnia intestiniformis but the conidia measure 
5-6*5 x 0*8—1 //m (Hillmann, 1936 : 80). If Gerber's figures are reliable, there can conse- 
quently be little doubt that his assertion was correct in that he was not dealing with the 
normal pycnidia of the host. The name must, however, remain of uncertain application until 
fresh material that might be referrable to Gerber's taxon is discovered, and the method of 
conidiogenesis determined. 

Libertiella xanthoriae Keissler, Annln naturh. Mus. Wien 38 : 166 (1925). 

Type: Sweden, Vestmanland, Vesteras, on Xanthoria fallax (Hepp) Arnold [?], 2 August 
1924, J. A. Nannfeldt(W 1925/10417— holotype!). 

The Xanthoria thalli appear to be quite healthy and are their normal colour, except 
for some deeper orange convex swellings (200-600 jum wide). In vertical section these 
were seen to comprise a very irregular compact conidiiferous structure and to produce 
numerous hyaline subcylindrical to rather ellipsoid conidia 3^1 x 1-2 jum which arose 
laterally on integrated doliiform conidiogenous cells. This area was not sharply delimited 
from the host tissues internally and the hyphae merged indistinguishably with the rest of the 
thallus, which included healthy algal cells in the immediate vicinity of the pycnidia. 

Pycnidia (spermogonia) are not common in X. fallax but were figured by Galloe 
(1948 : pi. 98 fig. 641) as deeper orange convex swellings superficially identical to 
those on Nannfeldt's specimen. As the conidiogenous cells are similar to those of 
other Xanthoria species (see e.g. Galloe, op. cit.) it is evident that Keissler's name 
was based on the normal pycnidia of the host; the species of Xanthoria may not have 
been X. fallax, however, as the conidia in that species are reported to be a rather 
more elongate (Moberg, 1977 : 13). 

Lichenophoma haematommatis Keissler, Hedwigia 50 : 296 (1911). 
(Fig.35A-B) 

Type: Austria, "in valle flumius **Enns , \ dicto "Gesause", ad Gesause-Eingang\ on Haematomma 
elatinum (Ach.) Massal. thallus, July 1910, AT. Keissler. 

The new genus Lichenophoma Keissler was introduced by Keissler (loc. cit.) for this 
single species. Unfortunately the type material could not be located in W (U. 
Passauer, in litt.) and, as noted by Sutton (1977: 113-114), the structure of the wall, 
conidiogenesis, and relationship of the sterile paraphyses-like filaments figured (Fig. 



78 D. L. HAWKSWORTH 

35b) to the conidiophores all require clarification before any firm opinion on the 
genus can be reached. The pycnidia were described as immersed, c. 120-150x60- 
80 /zm, with conidiogenous cells 12-15 x \ jum (!) forming oblong conidia which 
were hyaline and c. 5x2 pm; the sterile filaments, c. 60-70 x 1 //m, branched and 
approached the ostiole. I have not seen any fungus on Haematomma elatinum that 
could be identical with Keissler's taxon. There is a second report of this species by 
Keissler (1933 : 390) from a sterile lichen thallus on Quercus in France (Oise, Foret 
de Compiegne, au mont St Marc, November 1923, E. Cottereau) but that collection also has 
not been located. 

Lichenophoma opegraphae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 285 (1914). 

This taxon was based on two collections of Bouly de Lesdain from northern France: 
on Opegrapha sp. at Merckeghem, and on O. niveoatra (Borrer) Laundon (syn. O. 
subsiderella (Nyl.) Arnold) from Ghyvelde. The pycnidia were described as immersed, 
100-1 50 //m diam, with pseudoparenchymatous brownish walls composed of polyhedral 
cells 3-6 pm thick. Oblong conidia somewhat narrowed at the base, often unequal-sided, 
simple, hyaline, and 3*5-5 x 1*5-2 //m, were given as produced from simple conidiophores, 
10-22 x 1-5 jum. Sterile branched and anastomosing hyphae 2-2-5/zm thick formed 
k un reseau irregulier'. This taxon is not represented in the remnants of Vouaux's herbarium 
(Rondon, 1970) and as that of Bouly de Lesdain was destroyed in 1940 the application of this 
name must remain uncertain. It is, however, probable that Vouaux was dealing with the 
microconidial state of the host which forms unequal-sided conidia mainly 4-6(-7) x 
1-1 -5 pm. It is most unlikely that he had the fungus treated here as Laeviomyces 
opegraphae (p. 28) as that has much shorter conidiogenous cells, pale brown conidia with 
equal sides and a distinctly truncated base, and no network of hyphae in the pycnidial cavity. 

Microthelia solorinaria Lindsay, Q. Jlmicrosc. Sci. II, 9 : 350 (1869). 

Type: Ireland, Co. Kerry, Brandon Mountain, on Solorina crocea (L.) Ach., /. Carroll 
(E — holotype!). 
Diplodina solorinaria (Lindsay) Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 289 (1914). 

In the original place of publication of this name, Lindsay (loc. cit.) stated merely 
'M. solorinaria, Linds., Spermog., 175. On thallus of Solorina crocea. Pycnidia only.' 
Although no description was provided here, as the earlier publication of Lindsay 
(1859 : 175) is cited, and a description was given there, the name must be accepted as 
validly published. Lindsay (1859 : 175) cited two specimens 'Brandon Mountain, Kerry, 
Ireland, CARROLL; Wicklow, D. MOORE, in Herb. Carroll' but only one of these 
(unspecified) had the fungus concerned. Both were thought to be lost (Hawksworth, 
1980/? : 174) but the former was recently re-discovered misfiled under Pleospora hookeri 
(Borrer) Keissler in E; as this has a fungus conforming to Lindsay's description and drawings 
it must be the holotype. This collection supports what is certainly Rhagadostoma lichenicola 
(de Not.) Keissler; although no intact asci were seen the ascospores were typical for that 
species and 'Munk-pores' were present on the peridial cells. Dacampiosphaeria rivana (de 
Not.) D. Hawksw. is also present on the specimen but does not predominate nor conform to 
Lindsay's account. The name M. solorinaria is consequently correctly interpreted as a later 
synonym ofR. lichenicola. 

Phaeoantenariella lichenicola Cavalcante, Publcoes Inst. Micol. Recife 633 : 3 (1969). 

Type: Brazil, Maranhao, alto Turi, on Mazosia melanophthalma (Mull. Arg.) R. Sant. on 
indet. leaves, 27 December 1 967, J. de Anchieta (URM 80709 n.v.). 

The monotypic new genus Phaeoantenariella Cavalcante was introduced for a 
fungus with broad, torulose, dark brown hyphae spreading over the surface of the 
host, and which was reported to produce minute intercalary pycnidia, only 1-3 times 
the width of the hyphae, and conidia which were brown, simple, and 2-4 x L5-2//m. A 



LICHENICOLOUS COELOMYCETES 79 

specimen received from Recife and labelled 'tipo' from the type locality was examined but 
this bore the date 8 April 1 967 and was numbered URM 7 1 963 (exs. 4 1 664!), contrary to the 
type description. This material had, however, been studied by Cavalcante in July 1968, prior 
to the publication of the name, and supported hyphae certainly identical to those figured by 
Cavalcante (loc. ciL: 4-5). The 'pycnidia' described appeared to be only compact dividing 
regions of cells, and the 'conidia' must be assumed to be merely cells that had become 
dissociated by pressure during slide preparation. 

This name is consequently rejected as based on sterile mycelium. 

Phoma abietinae Vouaux, Bull, trimest. Soc. rnycol. Fr. 30 : 282 (1914). 

Type: Switzerland, k ad truncos abietinos in m. Gurnigel', L. E. Schaerer [Lich. Exs. no. 
534] (E— holotype!). 

Exsiccata: Schaerer, Lich. Exs. no. 534 (E!; sub Lecidea leucocephala C. patelliformis). 

Vouaux (1914 : 282) based his account of Phoma abietinae on Lindsay's (1872 : 259) 
description of pycnidia in the above exsiccatum; he did not indicate that he had studied any 
examples for himself of this number. Pycnidia are now rather sparse in the holotype, 
scattered over the thallus of Lecanactis abietina (Ach.) Korber, which has numerous 
apothecia, and agree in all details with the pycnidial state of this lichen (see Coppins & 
James, 1979); the C+ red reaction obtained from the pruina covering the aging pycnidia, the 
shape of the pycnidia, and size of the conidia, all show that this is not L. subabietina Coppins 
& P. James, a species with C— pycnidial pruina not known to form apothecia. The name 
Phoma abietinae should consequently be added to the synonymy of Lecanactis abietina, and 
excluded from the lichenicolous fungi. 

Phoma alectoriae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 197 (1914). 

Type: Switzerland, k ad Abietum truncos in m. Gurnigel', on Bryoria capillaris (Ach.) 
Brodo & D. Hawksw., L. E. Schaerer [Lich. Exs. no. 496] (E — lectotype!). 

Exsiccata: Schaerer, Lich. Exs. no. 496 (E!; sub Parmeliajubata S. carta). 

lcon.es: Lindsay, Trans. R. Soc. Edinb. 22 : pi. IV figs. 17-18(1859). 

Vouaux's (1914: 197) tentative introduction of this name for a species he considered allied 
to Phoma usneae (see p. 57) was based entirely on Lindsay's (1859 : 134) description and 
illustrations. The flattened blackened warts mentioned by Lindsay now occur rather sparsely 
on the lectotype and in none could any pycnidia or conidia be found. These warts do, how- 
ever, closely resemble the young ascocarps of Phacopsis huuskonenii Rasanan, which is 
well-known on the same host (Hawksworth, 1978a). The 'sterigmata' figured recall the 
paraphyses of P. huuskonenii, and furthermore the spore size given (15-23 x 3 jum) is 
reminiscent of that of this same species (( 1 2— ) 1 4—1 6(— 1 8) x 2-3 '5 jum, fide Hawksworth, loc. 
cit.), although the conidia in Lindsay's figure do not belong here. Lindsay's figure 
was, however, also based on Schaerer's Lich. Exs. no. 392 (sub Parmelia mollis y. 
divaricata; E!), but this packet no longer contains any intermixed Bryoria. 

This name, which in any case might be interpreted as not validly published under 
Art. 34, is considered to have been based on Phacopsis huuskonenii and some other 
fungus. As neither of these elements constitutes a satisfactory type, Vouaux's name 
must be rejected. 

Phoma arachnoidea Gerber, Arch. Protistenk. 74 : 483 (1931 ). 

Type: Austria, Solden, on Hvpogymnia intestiniformis (Vill.) Rasanen (syn. H. encausta 
(Sm.) W.Watson), W.Zopf. 

No material under this name could be located in B (B. Hein, in litt.) or DR (see 
p. 88). The pycnidia were described as about half-immersed and producing ellipsoid, 
hyaline, and often pointed conidia 3-5^-25 x L75-2-5//m, formed apically on filiform, 
non-septate, smooth conidiophores, measuring 12x2 , 5/zm. If Gerber's measurements are 



80 D. L. HAWKSWORTH 

correct and his description of the conidiophores accurate, he must have been dealing with a 
lichenicolous fungus and not the normal pycnidia of the host (see p. 77). Gerber's name 
consequently remains of uncertain application, but the long conidiophores certainly exclude 
it from Phoma. 

Phoma biformis Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 282 (1914). 

Type: France. Moeres, on Anisomeridium biforme (Borrer) R. C. Harris thallus on Salix 
alba wood, M. Boulyde Lesdain. 

No material of this taxon is present amongst the remnants of Vouaux's herbarium 
(Rondon, 1970), and that of Bouly de Lesdain was destroyed in 1940. The pycnidia 
were described as 50-140 pm wide, and as producing ovoid to globose conidia 
1-1*75 x 1-1-5 jim on conidiogenous cells 5-\2 pm long. Normal pycnidia of this 
lichen are not uncommon but may be of two types: microconidia which are orbicular 
and 2-3 pm, or macroconidia which are orbicular to elliptical or ovate and 3-5 pm 
diam or 3'5-5 x 2-5—3 pm (Harris, 1975:104). Vouaux's measurements suggest that 
he was not therefore dealing with the pycnidia of the reported host, but it is quite 
possible mistakes were made in measuring such small conidia. The name remains of 
uncertain application. 

Phoma caperatae Vouaux, in Bouly de Lesdain, Bull. Soc. hot. Fr. 59 : 16(1912). 

Type: France, Versailles, Fausses-Reposes wood, on Parmelia caperata (L.) Ach. on oak, 
M. Bouly de Lesdain. 
Phoma physciicola var. caperatae (Vouaux) Keissler, Beih. Bot. Zbl. II. 37 : 273 (1920). 

This species is not represented in the remnants of Vouaux's herbarium (Rondon, 
1970), and that of Bouly de Lesdain was destroyed in 1940. The description of the 
pycnidia as superfical, black, subglobose and 0* 1-0*25 mm diam is suggestive of 
Vouauxiomyces truncatus (see p. 70), but the ovoid conidia were described as 'basi 
saepe angustatis atque acutiusculis' — most inappropriate for that species. The conidia were 
given as 3-6 x 2*5-3 pm, and the conidiogenous cells as 3-4 x 2 pm. Bouly de Lesdain (loc. 
cit.) mentions a second collection on the sterile thallus of a Calicium species on oak at 
Rodez, Aveyron; these dimensions are suggestive of the normal conidia of Calicium rather 
than any lichenicolous fungus known from this host. The collection on Parmelia at least 
must have been a lichenicolous fungus, but its position remains uncertain in the absence of 
the original material. 

Phoma curvispora Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 281 (1914). 

In the original description of this species Vouaux cited three specimens, all collected by 
Bouly de Lesdain: on Lecanora saligna (Schrader) Zahlbr. from Versailles, on Lecania sp. 
from Zuydcoote, and on Lecidella elaeochroma (Ach.) M. Choisy from Ghyvelde. Unfortu- 
nately Bouly de Lesdain's herbarium was destroyed in 1940, and this taxon is not represented 
amongst the remnants of Vouaux's herbarium (Rondon, 1970). Keissler (1930 : 547) states 
that he saw the 'Originalexamplar' himself, and that it represents only pycnidia of the lichen 
in question but does not indicate which of the three original specimens he was able to see. 
The description of the conidia as 'tres courbes, souvent en demi-cercle', and 8-1 5 x 2-3 pm, 
leaves little doubt that the name Phoma curvispora was partly based on the pycnidia of 
Lecanora saligna. 

Phoma epiparmelia Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 282 (1914). 
Type: Yugoslavia, Nadin, Gradina, on Peltigera pusilla Korber, Baumgartner. 

This fungus was described as not damaging the host thallus and comprising erumpent, 
black, globose pycnidia 60-1 30 //m diam, with pseudoparenchymatous walls made up of 
brown cells 7-10 x 4-6 pm. Unbranched conidiophores, simple or rarely 1 -septate, sinuous, 
and 6-10 x L5-2 pm gave rise to rather irregularly shaped ellipsoid to frequently unequal- 



LICHENICOLOUS COELOMYCETES 81 

sided simple hyaline conidia measuring 3 -5-4- 5 x 2-3 //m. Unfortunately the original 
material could not be located in W (H. Riedl, in litt.), and is not present amongst the 
remnants of Vouaux's herbarium (Rondon, 1970). The long conidiogenous cells, smaller 
pycnidia, and lack of damage to the host, separate this taxon from Phoma peltigerae (see 
p. 54). In the absence of the original material the name must be treated as of uncertain 
application until a fungus conforming to Vouaux's description is re-discovered. 

Phoma epiphyscia Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 197 (1914). 

Type: France, Ghyvelde, on Phaeophyscia orbicularis (Necker) Moberg and Xanthoria 
parietina (L.) Th. Fr., M. Bouly de Lesdain. 

This taxon is not represented amongst the remnants of Vouaux's herbarium (Rondon, 
1970), and as that of Bouly de Lesdain was destroyed in 1940, the type material is presumed 
to be lost. The species was described as having immersed black pycnidia 80-1 50 //m, with 
pseudoparenchymatous walls, and conidia hyaline, simple, 2-guttulate, 4-6 x 2-3 //m, 
formed from short conidiogenous cells. The conidial dimensions given are too large for 
Phaeophyscia orbicularis and Xanthoria parietina. Vouaux (loc. cit.) suggested that it might 
represent an anamorph of the species now called Arthonia epiphyscia Nyl.; I have not seen 
pycnidia associated with A. epiphyscia, but they may occur (see p. 72). A further possibility 
is that Vouaux was dealing with the anamorph of Guignardia olivieri (see p. 73), as 
suggested by Hawksworth (1975/?: 191). 

Phoma fusispora Vouaux, in Bouly de Lesdain, Bull. Soc. bot. Fr. 59:215 (1912); 
nom. illegit. (Art. 64). 
Non Phoma fusispora Rostrup, Bot. Faeroes 1 : 3 1 3 ( 1 90 1 ). 

This species was originally described from specimens of Lecanora saligna (Schrader) 
Zahlbr. and L. dispersa (Pers. ex Sommerf.) Nyl. collected by Bouly de Lesdain k aux environs 
de Dunkerque'. It was also mentioned by Bouly de Lesdain (loc. cit.) as on Buellia punctata 
(Hoffm.) Massal. Vouaux (1914 : 196) added Bacidia spododes (Nyl.) Zahlbr.* as a host and 
Bouly de Lesdain (1914 : 165) Opegrapha niveoatra (Borrer) Laundon. Bouly de Lesdain's 
herbarium was destroyed in 1940, and this species is not represented in the remnants of 
Vouaux's herbarium (Rondon, 1970). However, Keissler (1930 : 547) reports that he was 
sent the 'Originalexemplar' by Bouly de Lesdain and that the name was based on the 
pycnidia of the host, but does not indicate which host he received. The conidia were origin- 
ally described as Tusiformibus, paulum arcuatis, continuis, hyalinis' and 7-8 x 2 jum, 
characters seen in numerous lichen pycnidia. In the absence of material some doubt must 
remain, but the evidence suggests that this name was based on the pycnidia of several 
different lichens. 

Phoma glaucella Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 198 (1914). 

Three French collections were mentioned in the original account of this species: on 
Bacidia inundata (Fr.) Korber at Malo-Terminus, on Lecanora sp. at Ghyvelde, and 
on L. piniperda Korber (syn. L. glaucella (Flotow) Nyl.) at Meyrueis, Lozere; the first 
two collections were made by Bouly de Lesdain and the last by Marc. The species was 
described as having mainly immersed, black pycnidia 60-140 //m diam, simple or 
rarely bifurcate conidiogenous cells 5-10 x 1 -5-2 //m, and ellipsoid, hyaline conidia 
L5-3 x 1-1 -5 jum. As no material is present in the remnants of Vouaux's herbarium 
(Rondon, 1970), and that of Bouly de Lesdain was destroyed in 1940, the application 
of this name remains uncertain. 

Phoma lecanorae Vouaux, in Bouly de Lesdain, Rech. Lich. Dunk.: 277 (1910). 

Type: France, Dunkerque, dunes pres le Phare, l sur Lecanora umbrina var.\ 18 June 
1904, M. Bouly de Lesdain (herb. Vouaux — syntype!). 

* = Micarea nitschkeana (Lahm ex Rabenh.) Harm. 



82 D. L. HAWKSWORTH 

Two specimens were mentioned in the original description of this taxon, the above 
and one from Malo-Terminus reported as on Rinodina exigua (Ach.) Gray; this latter 
collection is not represented in Vouaux's herbarium (Rondon, 1970), and as Bouly de 
Lesdain's herbarium was also destroyed, must be presumed lost. The collection from 
near Le Phare comprises a blackish-brown thallus on flint over which the pycnidia 
are evenly scattered; fruits of Lecanora polytropa are scattered amongst the dark 
thallus and were presumably mistaken for L. dispersa (Pers.) Sommerf. (syn. L. 
umbrina Nyl.) by Vouaux, but appear to be quite unrelated to the pycnidia. The 
evenly distributed pycnidia and healthy thallus leave no doubt that this material is 
the pycnidial state of a lichenized species; the thallus colour and short conidia 
(3-4 x 1-1*5 jum) are suggestive ofOpegrapha lithyrga Ach. 

Bouly de Lesdain (1914) and Vouaux (1914) reported eleven further hosts from a 
wide range of genera (interestingly, these included two Opegrapha species); these 
cannot be said to have comprised only lichen pycnidia in the absence of the 
collections cited. 

Phoma lichenis Pass., Atti Accad. naz. LinceiRc. 7 : 48 (1891). 

Type: Italy, prov. Parma, Vigheffio, Frassino, 'sul tallo di un Lichene sterile, fosse Parmelia 
pulverulenta' [Physconia pulverulacea Moberg], G. Passerini. 

The type material could not be located in either PAD (S. Chiesa, in litt.) or 
PARMA (F. Lona, in litt.). The black pycnidia were described as having mem- 
branaceous pseudoparenchymatous walls and producing narrow bacilliform conidia 
5 pm long. Conidia 4-7 x 1*5 pm are, however, characteristic of the pycnidia in both 
Physcia (Schreber) Michaux and Physconia Poelt (Moberg, 1977), and it therefore 
seems probable that this name was merely based on the pycnidia of the host. If this is 
so, as Art. 59 does not apply to lichen-forming fungi, Passerini's name may provide 
an earlier epithet for Physconia pulverulacea. 

There are reports of this taxon from a wide range of hosts, some of which may refer 
to true lichenicolous fungi. Keissler (1930:541-2) mentions alsp Cladonia rangi- 
ferina (L.) Wigg., Lecanora pallida (Schreber) Rabenh., Physcia stellaris (L.) Nyl., 
and Xanthoria parietina (L.) Th. Fr.; Keissler (1933 : 386) adds Cladonia rangiformis 
Hoffm. and Parmelia laevior Nyl. 

Phoma lichenis f. immersa B. de Lesd., Rech. Lich. Dunk. : 277 (1910). 

Type: France, Dunkirk, on Hypogymnia physodes (L.) Nyl., M. Bouly de Lesdain. 

As Bouly de Lesdain's herbarium was destroyed in 1940, the type collection is 
presumably lost. The only description provided was 'Les peritheces sont enfonces au 
lieu d'etre superficial'. Assuming that the conidia were roughly the same size as those 
given for Phoma lichenis, there can be little doubt that this form was based on normal 
pycnidia of the host. 

Phoma verrucariae Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 28 1 (19 14). 
Type: France, Bergues, on Verrucaria sp. [?], M. Bouly de Lesdain. 

This species was described as producing 2-3 pycnidia per areole of the sterile host, 
the pycnidia as immersed to almost superfical, black, 60-1 10 jum diam, with conidiophores 
10-20/zm long, and conidia which were cylindrical, attenuated at the ends, slightly or 
strongly curved, hyaline, simple, and 12-14 x 1-8-2'5/im. The host may not have been a 
Verrucaria as it was sterile and so could not have been checked microscopically; the name 
could therefore have been based on pycnidia of the 'host' and not a lichenicolous fungus. As 
this taxon is not represented in the remnants of Vouaux's herbarium (Rondon, 1970), and 
that of Bouly de Lesdain was destroyed in 1940, the application of this name remains 
uncertain. 



LICHENICOLOUS COELOMYCETES 83 

Phoma versoniana Sacc, Michelia 2 : 342 (188 1); nom. inval. (Art. 32). 

Keissler (1930:582) lists the name as a possible synonym of the species now called 
Libertiella malmedyensis, but it does not appear to have been ever validly published 
and the only mention of it I have been able to locate is the reference cited above 
where the phrase 'Phomae Versonianae meae analogo videtur' appears. It is not even 
known whether P. versoniana was considered to be lichenicolous. 

Phyllosticta galligena F. Moreau, Bull Soc. bot. Fr. 98 : 102 (1 95 1). 

Type: India, Palni Hills, Shembaganur, on Parmelia perforata Taylor, 1933, G. Foreau. 

Icones: Moreau, Bull. Soc. bot. Fr. 98 : 101 (1951). 

The type material of this gall-forming species could not be located in PC (J. 
Mouchacca, in litt.), nor was it available from Dr C. Moreau. Convex galls 0*25—1 5 mm 
were formed each including several black pycnidia (to 150//m diam) forming simple, 
ellipsoid, hyaline conidia 1-5-3*5 x 1-2 jum. The conidiophores were described as 
unbranched and filiform, so it is clear that this species cannot be referred to Phoma (see 
p. 49). In the absence of further information it is impossible to suggest an appropriate genus 
for this species, but it should be possible to recognize it if re-collected and then re-assess its 
position. 

Phyllosticta lichenicola Allescher, Ber. bayer. hot. Ges. 4 : 32 (1896). 

Type: Germany, Miinchen, Ebenhausen, on Parmelia perlata (Huds.) Ach., 1 1 April 1895, J. N. 
Schnabl (M — holotype!). 

This fungus was originally described as forming in necrotic patches about 2 mm 
diam, and having globose, black pycnidia with conidia hyaline, oblong, biguttulate, 
and 2-\ x 0*5-1*5 jum. Keissler (1930:535) studied the type material superficially 
but made no microscopic preparation because it was so small; no later authors appear 
to have found any fungus matching Allescher's description exactly, so a careful study 
of this specimen was needed. Only a single infection spot 2 mm diam remained on the 
thallus, this was necrotic with a black margin about 125-200 jum wide and contained 
six small black immersed sporocarps. Three of these proved to be ascomata of a 
Polycoccum or Endococcus species (paraphyses doubtfully present and not clearly 
demonstrable) with asci about 30 x 7 jum and 1 -septate, pale brown, echinulate ascospores 
about 8x3 //m; the asci were 6-8 spored. A fourth was immature and without spores, and 
the fifth a pycnidium with subglobose to broadly ellipsoid, hyaline conidia 3*5-4*5 x 
2-3*5 //m; no conidiogenous cells could be found. The sixth sporocarp was left un- 
examined for future investigators. Representative slides from the sporocarps examined are 
preserved in IMI. 

In view of these observations, it is clear that Allescher's taxon was based on 
discordant elements, the two fungi found by me and a third, perhaps Phoma cytospora, with 
conidia approximating to the shape and width he indicated. This name cannot be satis- 
factorily typified by these disparate elements and must consequently be rejected. 

Pleurosticta lichenicola Petrak, Kryptog. Forsch. 2(2) : 190 (193 1). 

This new genus and species was introduced by Petrak (loc. cit.) for what he inter- 
preted as a lichenicolous coelomycete growing on Parmelia olivacea (L.) Ach. in 
Germany. The identity of this taxon has been investigated by Santesson (1949: 142) 
who found it to be based on the normal pycnidia of a different lichen, P. acetabulum 
(Necker) Duby. Unfortunately the type collection could not be re-located in W (H. 
Riedl, in litt.), but there seems to be no reason to question Santesson's interpretation. 

Pleurosticta Petrak must consequently be treated as a further synonym of Parmelia 
Ach. nom. cons. Further, as Art. 59 specifically excludes lichenized fungi, if the 



84 D. L. HAWKSWORTH 

group of brown Parmelia species for which Esslinger (1978) introduced the new genus 
Melanelia Essl. are regarded as generically distinct, then Pleurosticta would be an 
earlier name for that genus unless conserved, as proposed by Esslinger (1980). 

Pseudodiplodia lichenis Vouaux, in Bouly de Lesdain, Reck. Lick. Dunk., Suppl. : 167 
(1914). 

Type: France, Malo Terminus, on a telegraph pole, 28 August 1910, M. Bouly de Lesdain 
(herb. Vouaux — holotype!). 
Stagorwpsis lichenis (Vouaux) Keissler, Rabenh. Krypt.-Fl. 8 : 586 (1930). 

This species was described as a fungus occurring on Lecanora saligna (Schrader) 
Zahlbr. (syn. L. effusa (Hoffm.) Ach.), and has pale orange, cupuliform pycnidia 
1 00-200 pm wide evenly distributed over a greyish-green crustose thallus; the hyaline 
conidia are curved with rounded ends, 0-1 septate, and 8-10 x 2-A jum. No Lecanora 
apothecia were found on the specimen. The lack of damage to the host as well as the even 
distribution of the pycnidia on the thallus leave no doubt that this name was based on the 
pycnidial state of an unidentified lichenized fungus. The shape of the conidia recalls that 
characteristic of the L. dispersa (Pers.) Sommerf. group within Lecanora, but the orange 
pycnidia are atypical for that facies. 

Vouaux's name was transferred to Stagonopsis by Keissler (1930:586) primarily 
because the conidia could be 1 -septate. 

Pyrenochaeta collematis Vouaux, in Bouly de Lesdain, Rech. Lich. Dunk. : 275 (1910). 

Type: France, Dunkerque, Bray Dunes, on apothecia and thalli of Collema tenax (Swartz) 
Ach., M. Bouly de Lesdain. 

This fungus was originally described as forming black, almost superficial pycnidia 
100-1 50 //m diam, with 3-9 setae 40-60 x 3-5 jum arranged around the ostiole, and 
producing hyaline simple conidia 5-6 x 2 pm on conidiogenous cells 6-7xl*5/im. 
Vouaux (1912:185, 1914:288) suggested that it might represent an anamorph of 
Nectria tenacis (Vouaux) Vouaux, but it would be remarkable for any Nectria to have 
a coelomycete state; Pyrenochaeta de Not. species have teleomorphs in Herpotrichia 
Fuckel. As Bouly de Lesdain's herbarium was destroyed in 1940, and this taxon is not 
represented amongst the remnants of Vouaux's herbarium (Rondon, 1970), it has not 
been possible to examine any material of this taxon. According to Vouaux's descrip- 
tion this fungus cannot have been a Pyrenochaeta because the conidiogenous cells in 
this genus are arranged in long acropleurogenous chains (Schneider, 1979). 

Rhabdospora antarctica Speg., An. Mus. nac. B. Aires 20 : 390 (1910); as 4 n.f.' 

Type: Antarctic Islands, South Orkney Islands ['Nuevas Orcadas'], on Caloplaca regalis 
(Vainio) Zahlbr., February 1908, C. Spegazzini (LPS 1 1231— holotype!). 

The host, originally indicated to be a Teloschistes Norman species, proves to be the 
ubiquitous antarctic polymorphic Caloplaca regalis (syn. Polycauliona regalis (Vainio) 
Hue). In the original description of this fungus the 'perithecia' were given as black and 
75-90 pm diam, and the spores as hyaline, 1 -septate or simple, 'rotundato-subtruncatae' 
and 12-16 x 2-3 //m. The holotype is very small, the host fragment measuring only 
3x2 mm, but two sporocarps recalling Spegazzini's description were present; one was 
examined and found to be the pyrenomycete Polycoccum rugulosaria (Lindsay) D. Hawksw., 
which if confined to this host and described and illustrated elsewhere (Hawksworth in Pegler 
et al, 1980). A few of the normal pycnidia of the Caloplaca were also found but they form 
conidia quite unlike those described by Spegazzini (loc. cit.). It is consequently clear that 
Rhabdospora antarctica should be treated as a synonym of P. rugulosaria. That Spegazzini 
gave the spores as hyaline and sometimes simple, and also only 2-3 pm wide, 



LICHENICOLOUS COELOMYCETES 85 

indicates that he based his description on some immature ascospores. Neither asci 
nor conidiogenous structures were mentioned by Spegazzini. 

The name Rhabdospora antarctica was evidently missed by Vouaux (1914); as a 
result it seems to have been overlooked by subsequent students of the lichenicolous 
fungi (e.g. Clauzade & Roux, 1976). 

Rhabdospora lecanorae Vouaux, in Bouly de Lesdain, Reck. Lick. Dunk., Suppl. : 167 
(1914). 

Type: Spain, Catalogne, Hostaleto, nr Figueras, on Lecanora chlarona (Ach.) Nyl. on 
olives, 1 1 April 1905, F. Heribaud (herb. Vouaux — lectotype!). 

In Bouly de Lesdain's publication three collections were cited, on Opegrapha vulgata 
(Ach.) Ach. (syn. O. cinerea Chev.) from Dunkirk, on Micarea nitschkeana (syn. Bilimbia 
spododes (Nyl.) Arnold) thalli from Bergues, and on Lecidella elaeochroma (Ach.) M. Choisy 
(syn. Lecidea parasema auct.) from Ghyvelde. Vouaux's selection of the epithet 'lecanorae', 
however, implied that he had a fourth collection in mind when providing his description for 
Bouly de Lesdain, on Lecanora. Vouaux (1914 : 290) cites a collection on L. chlarona first, 
before the Bouly de Lesdain material, and this must be regarded as a syntype for this name. 
As only this latter collection is extant it is here designated as the lectotype. 

The type is in a poor condition and is overgrown by superficial moulds; in some 
apothecial margins acicular to arcuate or sigmoid hyaline conidia 20-30 x 1-1 -5 
(-2) jum, produced on pyriform conidiogenous cells 4-6 x 3-5 jum, were found. Identical 
conidia were also produced in black pycnidia directly on the thallus, something overlooked 
by Vouaux. These correspond to the pycnidia (spermogonia) usually produced by this 
lichen, and the name Rhabdospora lecanorae should consequently be treated as a synonym 
of Lecanora chlarona. 

Rhabdospora lesdainii Vouaux, Bull, trimest. Soc. mycol. Fr. 30 : 290 (1914). 

Type: France, Malo-Terminus, on Lecidella elaeochroma (Ach.) M. Choisy (syn. Lecidea 
parasema auct.) on Populus monilifera, M. Bouly de Lesdain. 

The type material is not amongst the remnants of Vouaux's herbarium (Rondon, 
1970), and as that of Bouly de Lesdain was destroyed in 1940, must be presumed lost. 
The fusiform, straight to slightly curved conidia were described as with 5-7 'plus ou 
moins distinctes' septa and given as 38-60 x 3^5 jum. The normal conidia of this 
species recall Vouaux's description of their shape but are not septate (only guttulate) 
and only 17-18 jum long (Galloe, 1927:92). The name is consequently of uncertain 
application. 

Rhabdospora thallicola Tassi, Bull. Lab. Orto Bot. Reale Univ. Siena 3 : 129 (1900). 
Type: Brazil, 'corteccie di Cinchona', September 1900, herb. F. Tassi (SIENA — holotype!). 

The type collection comprises two pieces of bark each covered with the creamy- 
white thallus of a sterile crustaceous lichen. The pycnidia are scattered rather regularly over 
the material and not associated with any signs of damage. The filiform conidia and arrange- 
ment of the conidiogenous cells is similar to that seen in the pycnidia of numerous lichens, 
particularly the Lecanora chlarotera Nyl. group. This name is consequently excluded as 
based on the pycnidia of an unknown lichen. 

Sphaeromma mazosiae Upadhyay, Publcoes Inst. Micol. Recife 402 : 4 (1964). 

Type: Brazil, Rondonia, Porto Velho, Perfil AR-4, on Mazosia melanophthalma (Mull. 
Arg.) R. Sant. on Orbigyna martiana, January 1962, O. M. Fonseca (URM 38672 
[19.945 p.p.]— holotype!). 



86 D. L. HAWKSWORTH 

The herbarium specimen URM 19.945 comprises twelve portions of leaf, mostly 
about 10-12 cm long, which support a wide range of lichens and associated lichenicolous 
fungi constituting Tungos do Brasil' nos. 38670-38682 (i.e. including the holotypes of both 
Sphaeromma mazosiae and Sporhaplus rondoniensis Upadhyay). Sphaeromma mazosiae 
and Sporhaplus rondoniensis were both described as having superficial, dark brown pycnidia 
(33^45 //m diam and 30-35 jum diam, respectively) with stout, dark brown setae (given as 
6—12*5 x 3*4 jum for each species); the Sphaeromma was stated to have 2-A septate conidia 
7-10*5 x 1-5-2 jum, and the Sporhaplus simple conidia 1 5—2 x 0*75—1 //m. Only after very 
extensive searching did I manage to locate any Mazosia thalli with setose sporocarps 
recalling those of the original description; several of these were squashed and found 
to contain pseudoparenchymatous tissue, young asci, or asci with 1 -septate hyaline 
immature spores about 8x4 jim. It seems probable that the ascomycete is identical to 
the monotypic Keratosphaera batistae Upadhyay, the holotype of which (URM 38673) 
should be on this specimen, although its spores were indicated to become 3-septate when 
mature and measure 6-10 x 2-2*5 jum, and no evidence for the existence of paraphyses 
figured by Upadhyay (loc. cit. : 10) was obtained. 

The host lichen was originally referred to Mazosia phyllosema (Nyl.) Zahlbr. by 
Upadhyay, but the thallus is not smooth but has the minute whitish verrucae 
characteristic of M. melanophthalma (Mull. Arg.) R. Sant., although they are rather 
poorly developed in this material. According to Santesson (1952 : 1 18), M. melanophthalma 
has macroconidia 5-8 x 2-3 jum and microconidia 2-3 x 1 /im; the correspondence be- 
tween the sizes of these and those given for Sphaeromma mazosiae and Sporhaplus 
rondoniensis is striking. In my opinion descriptions of both these fungi were probably based 
on the conidia of M. melanophthalma (the 'septation' of the larger conidia being due to 
guttulation) and sporocarps of the Keratosphaera. These two names cannot consequently be 
satisfactorily typified and must be rejected. 



Sphaeronema lichenophilum Durieu & Mont., in Montagne, Syll. gen. crypt. : 249 (1856). 

Type: Algeria, *sur les troncs du vieux Chamaeropis, du Marabout du sommet du Bouzareah', 8 
April 1 844, M. C. Durieu de Maisonneuve (PC — holotype!). 

This name has generally been cited as first published in Botanique I. Cryptogamiae 
of Durieu's Exploration scientifique de I'Algerie, which is referred to with correct 
page numbers in Montagne's book cited above. However, it is now clear that the part 
of Durieu's work including Sphaeronema lichenophilum lay in proof for at least 13 
and perhaps as many as 23 years before they were finally published in October 1869 
(Hawksworth & Booth, 1974: 12); only the first parts of this volume were issued in 
1846-50, the dates usually given for S. lichenophilum. Montagne cited only the single 
collection detailed above, which is consequently the holotype; Durieu mentioned the 
same collection, but also a further one made on Chamaeropis in February 1840 
which is also now preserved in PC (!). 

Durieu gave the host as 'Lecideae canescentis aut Dirinae Ceratoniae\ but in the 
holotype, the 1840 specimen, and further two small undated specimens in PC (!), the 
pycnidia are distributed over a whitish-grey, thin, crustose thallus forming circular 
patches mainly 5-10 mm diam. Eleven fragments of bark comprise the holotype 
sheet, ten of which have pycnidia conforming to the original description, but the 
other is Diploicia canescens (Dickson) Massal. unaffected by any pycnidia. In vertical 
section the pycnidia were found to extend down through the thallus to the bark and this, 
together with their even distribution and no evidence of damage to the thallus, strongly 
suggest that they are the pycnidia of a lichenized fungus, and not an independent 
coelomycete invading a lichen. The pycnidia are thick-walled, dark brown and 
carbonaceous, perennial with new pycnidia arising within effete ones, phialides mainly 
about 10 x 1*5 //m, and numerous, minute, simple, cylindrical, guttulate conidia mainly 



LICHENICOLOUS COELOMYCETES 87 

5-6 x 2-3 jum. Its identity is uncertain, although the pycnidia suggest Arthoniales or 
Hysteriales; it is not Dirina ceratoniae Fr., the pycnidia of which were illustrated by Vobis & 
Hawksworth(1981). 

Sphaeronema lichenophilum was also very tentatively reported from a Parmelia 
collected in 1921 in Brazil (Sao Paulo, Premembe) by Keissler (1933 : 385), but as the 
material was stated to be in Bouly de Lesdain's herbarium it was presumably 
destroyed in 1940. 

Sphaeropsis cladoniae Ell. & Ev., /. Mycol. 5 : 149 (1889). 
Coniothyrium cladoniae (Ell. & Ev.) Sacc, Syll Fung. 10 : 268 (1892). 

Lichenoconium cladoniae (Ell. & Ev.) Petrak & H. Sydow, Beih. Rep. nov. Spec. Regni veg. 
42:433(1927). 

This name was based on a collection including an Abrothallus species and Lichenoconium 
usneae, but is considered correctly used for the former element (Hawksworth, 
1977: 192-193). 

Sphaeropsis scripta Schwein. ex Berk. & M. A. Curtis, J. Acad. nat. Sci. Philad. 
2:280(1853). 

Dothiorella erumpens Sacc, Svli Fung. 3:242 (1884); as '(Berk. & C.)\ nom. illegit. (Art. 
63.2). 

This fungus was listed as a lichenicolous species by Viegas (1961 : 869, as 'Dothiorella 
erupens" [sic]) but Berkeley & Curtis (loc. cit., pi. 25 fig. 3a-c) illustrated a black, stromatic, 
pulvinate, multilocular conidioma bursting through the bark from below, and were evidently 
not dealing with a lichenicolous fungus; the fungus was erumpent through unidentified 
lichens on the bark, not on the lichens themselves. The conidia of the fungus were broadly 
ellipsoid, hyaline, simple, with a short persistent pedicel. 

Spilomium epicladonia H. Olivier, Paras. Lick Fr., Suppl. : 22 (1907) [not seen]. 
Type: France, Herault, Mons-la-Frivola, on Cladonia sp., A. de Crozals. 

No type or authentic material of this taxon has been located. Vouaux (1914: 198) 
considered it might be identical to the species now called Bachmanniomyces uncialicola, but 
this seems unlikely to be correct (see p. 14). 

Sporhaplus rondoniensis Upadhyay, Publcoes Inst. Micol. Recife 402 : 7 (1964). 

Type: Brazil, Rondonia, Porto Vehlo, Perfil AR-4, on Mazosia melanophthalma (Mull. 
Arg.) R. Sant. on Orbigvna martiana, January 1962, O. M. Fonseca (URM 38674 [URM 
19.945 p.p.]— holotype!)." 

The holotype of this monotypic genus is present on the same collection as that of 
Sphaeromma mazosiae and some other fungi, and is discussed under S. mazosiae 
above (p. 86). 

Sporocadus lichenicola Corda, Icones Fung. 3 : 24 (1839). 

Type: Czechoslovakia, Prague, Fiirstl. Lobkowitzischen Garten, on Rosa sp., 1838, A. J. 
C. Corda (PRM 155664— holotype n.v.). 
Hendersonia lichenicola (Corda) Fr., Summ. veg. Scand. 2:416 (1849); as \Sommerfy . 
Seimatosporium lichenicola (Corda) Shoem. & E. Miiller, Can. J. Bot. 42 : 405 (1964). 

This taxon, the lectotype species of the generic name Sporocadus Corda (Hughes, 
1958:810), was thought by Corda to be associated with some unidentified crustose 
lichen because algal cells were associated with the pycnidia of the fungus below the 
surface of the host. The type material was examined by Hughes (loc. cit.) and his 
slides were also studied by Shoemaker & Miiller (1964); these, together with numerous later 
collections identical to Corda's species, demonstrate conclusively that this is not a 
lichenicolous fungus, but rather a common saprophyte of Rosa stems. The algae present in 



D. L. HAWKSWORTH 




Fig. 36 Verrucaster lichenicola. A, Infected podetium (x2). B, Gall with pycnidia (x 12). 
C, Conidiophores (x 1000). D, Conidiogenous cells (x 1000). E, Conidia (x 1000). Reproduced 
from Tobler(1913: 383). 



Corda's collection occurred only fortuitously. Corda's fungus is correctly placed in 
Seimatosporium and has an extensive synonymy (Sutton, 1975 : 136); it is the anamorph of 
Clathridium corticola (Fuckel) Shoem. & E. Mliller (syn. Griphosphaeria corticola (Fuckel) 
Hohnel) the connection between the two having been demonstrated by cultures of the 
ascospores giving rise to the Seimatosporium state (Shoemaker & Miiller, 1964). 

In accepting " Render sonia lichenicola" as a lichenicolous species, Clauzade & Roux, 
1976 : 89) were presumably simply following Keissler (1930 : 578) who listed it from several 
lichen hosts; these must be presumed to be cases where the fungus grew through overlying 
lichen thalli or otherwise they were misidentifications. 

Sporocadus rosicola Rabenh., Bot. Ztg6 : 294 (1848); as 'rosaecola\ 

Type: Germany, prope Leipzig, 'ad Rosarum ramulos', L. Rabenhorst [Fungi Exs. no. 66] 
(not seen). 

Rabenhorst's fungus occurred on Rosa branches and there can be little doubt that 
its usual treatment as a synonym of Seimatosporium lichenicola (Corda) Shoem. 
& E. Miiller is correct (Sutton, 1975:136). However, Saccardo (1884:439) stated 
that it had been found \ . . in apotheciis Lecideae luteolae [i.e. Bacidia rubella 
(Hoffm.) Massal.], Gand in Belgio'. This later report must be treated as a misidentification 
based on an unknown lichenicolous fungus. 

Verrucaster lichenicola Tobler, Abh. naturw. Ver. Bremen 21 : 384 (1913) ['1912']. 
(Fig. 36A-E) 
Type: Germany, Oldenburg, Kehnmoor, on Cladonia bacillaris auct., H. Sandstede. 

This new genus and species was introduced by Tobler (1913) for a fungus forming 
gall-like growths on podetia of the host. The pycnidia were described as superficial, 
and opening irregularly, waxy, forming simple, hyaline, narrowly ellipsoid 2-guttulate 
conidia 3-6-7'6 x 0-8-1-6 jum from elongate conidiogenous cells apparently arranged on 
short conidiophores (Fig. 36C-D). The material on which this taxon was based has not been 
located in B (B. Hein, in litt.) or MSTR (B. Gries, in litt.); cryptogam collections in BRM are 
currently unavailable during re-housing (H. Kuhbier, in litt.), and any at DR would have 
been destroyed in 1945 (F. Mattick, in litt.). 



LICHENICOLOUS COELOMYCETES 89 

The galls produced by this fungus recall those of Epicladonia sandstedei (see 
p. 16), but the waxy, superficial pycnidia, and the simple and much narrower 
conidia, make it unlikely to belong there. Bachmanniomyces uncialicola is easily 
excluded as a possibility on the basis of conidial shape (p. 1 1), and E. simplex (p. 19) by the 
much shorter and narrower conidia. 

As I have seen no fungus recalling Tobler's description in all details, the applica- 
tion of his name must currently remain uncertain. However, the possibility that 
Tobler was dealing with an Epicladonia species cannot be entirely discounted. 



Acknowledgements 

Without the important studies of Dr B. C. Sutton on generic names and generic concepts in the 
Coelomycetes as a whole the present investigation would have been impossible. I have also 
benefited from his comments as to the placement of various taxa treated in this work, but stress 
that the decisions taken are my own. I have also continued to receive a steady stream of enig- 
matic specimens from Mr B. J. Coppins, which not only led to the discovery of some hitherto 
unrecognized taxa, but further enabled species concepts to be clarified through the study of 
more than single collections. 

Any revision of this type is heavily dependent on the availability of material from other herbaria. I 
am indebted to the curators and directors of all herbaria cited in the text (see p. 5) 
for enabling me to study collections in their care, and further to the following for assistance in 
trying to locate particular specimens: T. Ahti, S. Chiesa, B. J. Coppins, B. Gries, M. Guerlesquin, B. 
Hein, H. Hertel, H. Kuhbier, F. Lona, R. Moberg, J. Mouchacca, U. Passauer, H. Riedl, Y. Rondon, L. 
Tibell and O. Vitikainen. 

Miss F. J. Walker was of particular help in tracking down elusive collections at the British 
Museum (Natural History) and obtaining copies of some papers. Most of the sections illustrated were 
cut by Mrs C. Beer, and Mr D. W. Fry prepared the habit photographs included. 

Mr P. W. James and J. R. Laundon are thanked for their continuing interest in my studies on 
lichenicolous fungi and for their assistance in the publication of the present revision. 

References 

Bachmann, E. 1927a. Die Pilzgallen einigerCladonien. Arch. Protistenk. 57 : 58-84. 

\921b. Die Pilzgallen einigerCladonien. II. Arch. Protistenk. 59 : 373-416. 

1928a. Die Pilzgallen einigerCladonien. III. Arch. Protistenk. 62 : 261-306. 

1 928/7. Die Pilzgallen einiger Cladonien. IV. Arch. Protistenk. 64 : 109-1 5 1 . 

1929. Pilz-, Tier- und Scheingallen auf Flechten. Arch. Protistenk. 66 : 459-5 14. 

Batista, A. C. 1961. Um pugilo de generos novos de liquens imperfeitos. Publcoes Inst. Micol. 

Recife320:\-3l. 
Bouly de Lesdain, M. 1910. Recherches sur les Lichens des Environs de Dunkerque. Dunkirk. 

1914. Recherches sur les Lichens des Environs de Dunkerque. l er Supplement. Dunkirk. 

Cavalcante, W. A., Cavalcante, A. A. S. A. S. & Leal, F. B. 1972. Coletanea de liquens 

imperfeitos. Publcoes Inst. Micol. Recife 647 : 1-46. 
Christiansen, M. S. 1980. Lichenoconium erodens and some other fungi parasitic on Lecanora 

conizaeoides. Lichenologist 12 : 149-1 5 1 . 
Clauzade, G. & Roux, C. 1976. Les Champignons Lichenicoles non Lichenises. Montpellier. 
Clements, F. E. & Shear, C. L. 1931. The Genera of Fungi. New York. 
Coppins, B. J. & James, P. W. 1979. New or interesting British lichens IV. Lichenologist 

11: 139-179. 
Eigler, G. 1969. Studien zur Gliederung der Flechtengattung Lecanora. Dissnes bot, Lehre 

4:i-iii, 1-195. 
Esslinger, T. L. 1978. A new status for the brown Parmeliae. Mycotaxon 1 : 45-54. 

1980. Proposal to conserve Melanelia Essl. over Pleurosticta Petrak (Lichens). Taxon 29 : 692. 

Farr, E. R., Leussink, J. A. & Stafleu, F. A. (Eds). 1979. Index nominum Genericorum 

(Plantarum). [Regnum veg. 100-102.] 3 vols. Utrecht. 
Filson, R. B. 1978. A revision of the genus Heterodea Nyl. Lichenologist 10 : 1 3-25. 
Folan, A. C. M. & Mitchell, M. E. 1970. The lichens and lichen parasites of Derryclare Wood, 

Connemara. Proc. R. Ir. Acad. (B)70 : 163-170. 



90 D. L. HAWKSWORTH 

Fuckel, [K. W. G.| L. 1870. Symbolae mycologicae. Jb. nassau. Ver. Naturk. 23/24 : 1-459. 

Galloe. O. 1927-72. Natural History of the Danish Lichens. 10 vols. Copenhagen. 

Gerber, K. 1931. Die epiphytische Verbreitung von 'Flechtenparasiten'. Arch. Protistenk. 

74:471-489. 
Gliick, H. 1899. Entwurf zu einer vergleichenden Morphologie der Flechten-Spermogonien. 

Verh. naturh.-med. Ver. Heidelb. 11,6(2) : 81-216. 
Grummann, V. J. 1960. Die Cecidien auf Lichenen. Bot. Jb. 80 : 101-144. 

1969. Alte und neue Halbflechten. Ein neuer Flechtenparasit. Placynthium asperellum 

neu fur Mitteleuropa. Sydowia 22 : 216-224. 

Harmand, J. 1907. Lichens de France, fasc. 3. Paris. 

Harris, R. C. 1975. A taxonomic revision of the genus Arthopyrenia Massal. s. lat. (Ascomycetes) in 

North America. Ph.D. thesis, Michigan State University. 
Hawksworth, D. L. 1975a. A revision of lichenicolous fungi accepted by Keissler in Coniothecium. 

Trans. Br. mycol Soc. 65 : 219-238. 
1975/7. Notes on British lichenicolous fungi, I. Kew Bull. 30 : 183-203. 

1976. New and interesting microfungi from Slapton, South Devonshire: Deuteromycotina III. 

Trans. Br. mycol. Soc. 67 : 5 1-59. 

1977. Taxonomic and biological observations on the genus Lichenoconium (Sphaeropsi- 



dales). Persoonia9: 159-198. 

— 1978a. Notes on British lichenicolous fungi: II. Notes R. bot. GdnEdinb.36 : 181-197. 

— 1978/). The taxonomy of lichen-forming fungi: reflections on some fundamental problems. In 
Essays in Plant Taxonomy (H. E. Street, Ed.) : 21 1-243. London, New York & San Francisco. 

— 1979. The lichenicolous Hyphomycetes. Bull. Br. Mus. nat. Hist. (Bot.) 6 : 183-300. 

— 1980a. Notes on some fungi occurring on Peltigera, with a key to accepted species. Trans. 



Br. mycol. Soc. 74 : 363-386. 

— 1980/7. Notes on British lichenicolous fungi: III. Notes R. bot. Gdn Edinb. 38 : 165-183. 

— 1980c. Lichens of the South Devon coastal schists. Fid Studies 5 : 195-227. 

— & Booth, C. 1974. A revision of the genus Zopfia Rabenh. Mycol. Pap. 135 : 1-38. 
1976. Some observations on Nectria heterospora. Mycologia 68 : 195-200. 



Coppins, B. J. & James, P. W. 1980 ['1979']. Blarneya, a lichenized hyphomycete from 

southern Ireland. Bot. J. Linn. Soc. 79 : 357-367. 

& Dyko, B. J. 1979. Lichenodiplis and Vouauxiomyces: two new genera of lichenicolous 

Coelomycetes. Lichenologist 11 : 51-61. 

& Minter, D. W. 1980. New and interesting microfungi from the 1978 Exeter Foray. 

Trans. Br. mycol. Soc. 74 : 561-511. 

& Punithalingam, E. 1973. New and interesting microfungi from Slapton, South Devonshire: 



Deuteromycotina. Trans. Br. mycol. Soc. 61 : 57-69. 
Henssen, A. 1977. The genus Zahlbrucknerella. Lichenologist 9 : 1 7-46. 
Hertel, H. 1971. Uber holarktische Krustenflechten aus den venezuelanischen Anden. Willdenowia 

6:225-272. 
Hillmann, J. 1936. Parmeliaceae. Rabenh. Krypt.-Fl. 9, 5(3) : 1-309, 1-10. 
Hohnel, F. 1910. Fragmente zur Mykologie (XI. Mitteilung, Nr. 527 bis 573). Sber. Akad. 

Wiss. Wien 1, 119:617-679. 
Holmgren, P. K. & Keuken, W. 1974. Index Herbariorum. Part I. The Herbaria of the World. Ed. 6. 

[Re gnu m veg. 92.] Utrecht. 
Hughes, S. J. 1958. Revisiones Hyphomycetum aliquot cum appendice de nominibus rejiciendis. Can. 

J. Bot. 36 : 727-836. 
Keissler, K. von 1910. Einige bemerkenswerte Flechtenparasiten aus dem Pinzgau in Salzburg. 

6st.bot.Z. 60: 55-61. 

1911. Zwei neue Flechtenparasiten. Hedwigia 50 : 294-298. 

1927. Systematische Untersuchungen uber Flechtenparasiten und lichenoide Pilze. (V. 

Teil, Nr. 41-50). Annln naturh. Mus. WienAX : 157-170. 

1930. Die Flechtenparasiten. Rabenh. Krypt.-Fl. 8 : i-ix, 1-712. 

- 1933. Zusammenstellungeiniger interessanter Flechtenparasiten. Beih. bot. Zbl. 50 : 380-394. 
1958-60. Usneaceae. Rabenh. Krypt.-Fl. 9, 5(4) : i-ix, 1-755. 



Kendrick, [W.J B. 1980. The generic concept in Hyphomycetes — a reappraisal. Mvcotaxon 

11 : 339-364. 
Kotte, J. 1909. Einige neue Falle von Nebensymbiose (Parasymbiose). Zentbl. Bakt. ParasitKde II, 

24:74-93. 



LICHENICOLOUS COELOMYCETES 91 

Lindsay, W. L. 1857. Monograph of the genus Abrothallus (De Notaris and Tulasne emend.). 

Q. J I microsc. Sci. 5 : 27-63. 
1859. Memoir on the spermogones and pycnides of filamentous, fruticulose, and foliaceous 

lichens. Trans. R. Soc. Edinb. 22 : 101-303. 

1 869a. Observations on new lichenicolous micro-fungi. Trans. R. Soc. Edinb. 25 : 5 1 3-555. 

1869/). Enumeration of micro-lichen parasites on other lichens. Q. Jl microsc. Sci. II, 



9:49-57, 135-146,342-358. 

— 1871. Observations on lichenicolous microparasites. Q. Jl microsc. Sci. II, 11 : 28^42. 

1872. Memoir on the spermogones and pycnides of crustaceous lichens. Trans. Linn. Soc. 



Lond.28: 189-31 
Malloch, D. 1979. Plectomycetes and their anamorphs. In The Whole Fungus ([W.] B. Kendrick, Ed.) 

1 : 153-165. Ottawa. 
Mameli-Calvino, E. 1930. Ricerche su una forma singolare di deutoerohchene: Chlorocyphella sub- 

tropica Speg. Nuovo G. bot. ital. II, 37 : 369-379. 
Michaelides, J., Hunter, L., Kendrick, [W.J B. & Nag Raj, T. R. 1979. Icones generum 

coelomycetum. Supplementum. Synoptic key to 200 genera of Coelomycetes. University of 

Waterloo i$\o\)2§\ 1-41. 
Moberg, R. 1977. The lichen genus Physcia and allied genera in Fennoscandia. Symb. bot. 

upsal. 22(1): 1-108. 
Moreau, F. 1951. Sur une Dothideacee parasite du lichen Anaptychia speciosa (Wulfen) Massalongo. 

Bull. Soc. bot. Fr. 98 : 208-209. 
Morgan- Jones, G. 1971. Conidium ontogeny in Coelomycetes. III. Meristem thalloconidia. 

Can. J. Bot. 49: 1939-1940. 
Mttller [Argoviensisj, J. 1881. Lichenologische Beitrage XII. Flora, Jena 64 : 81-88, 100-1 12. 

1891. Lichenologische Beitrage XXXII. Flora, Jena 73 : 187-202. 

Nieuwland, J. A. 1916. Critical notes on new and old genera of plants. — VIII. Am. Midi nat. 

4:379-386. 
Nordin, I. 1964. Abrothallus suecicus, a common lichenicolous fungus. Svensk bot. Tidskr. 

58 : 225-232. 
Nylander, W. 1856. Prodromus lichenographiae Galliae et Algeriae. Actes Soc. linn. Bordeaux 

21:249-461. 

1858. Enumeration generale de lichens. Mem. Soc. Imp. Sci. nat. Cherbourg 5 : 85-146. 

Pegler, D. N., Spooner, B. M. & Smith, R. I. L. 1980. Higher fungi of Antarctica, the 

subantarctic zone and Falkland islands. KewBull. 35 : 499-562. 
Petrak, F. & Sydow, H. 1935. Kritisch-systematische Originaluntersuchungen iiber Pyrenomyzeten, 

Sphaeropsideen und Melanconieen. VI. Annls mycol. 33 : 157-193. 
Poelt, J. 1980. Eine diozische Flechte. PL Syst. Evol. 135 : 8 1-87. 
Punithalingam, E. 1974. Studies on Sphaeropsidales in culture. II. Mycol. Pap. 136 : 1-63. 

1979. Graminicolous Ascochyta species. Mycol. Pap. 142 : 1-214. 

Rizzini, C. T. 1952. Lichenes in Horto Botanico Fluminis Januarii crescentes. I. Archos Jard. 

bot., Rio de J. 12: 189-202. 
Rondon, Y. 1970. L'herbier des champignons parasites des lichens de Tabbe Vouaux. Revue 

bryol. lichen. II, 36 : 737-745. 
Saccardo, P. A. 1884. Svlloge Fungorum, 3. Padua. 

Sandstede, H. 193 1 . Die Gattung Cladonia. Rabenh. Krypt.-Fl. 9, 4(2) : 1-53 1 . 
Santesson, R. 1949. Svampar som leva pa lavar. Svensk bot. Tidskr. 43 : 141-143. 
1952. Foliicolous lichens I. A revision of the obligately foliicolous lichenized fungi. 

Symb. bot. upsal. 12(1) : 1-590. 

1960. Lichenicolous fungi from northern Spain. Svensk bot. Tidskr. 54 : 499-522. 



Schaechtelin, J. & Werner, R.-G. 1926. Developpement et biologie de V Abrothallus parmeliarum 

Smlft. Bull, trimest. Soc. mycol. Fr. 42 : 233-243. 
1928. Un cas foudroyant de parasymbiose. Le Homostegia piggotii (Berk, et Br.) 

Karst., son developpement biologique et physiologique. Bull, trimest. Soc. mycol. Fr. 44 : 232-240. 
Schneider, R. 1979. Die Gattung Pvrenochaeta de Notaris. Mitt. biol. BundAnst Ld-u. Forstw. 

189: 1-73. 
Shoemaker, R. A. & Muller, E. 1964. Generic correlations and concepts: Clathridium 

{=Griphosphaeria) and Seimatosporium (=Sporocadus). Can. J. Bot. 42 : 403^07. 
Sutton, B. C. 1968. Typification of Dendrophoma and re-assessment of D. ohscurans. Trans. 

Br. mycol. Soc. 48: 611-616. 



92 D. L. HAWKSWORTH 

1973. Coelomycetes. In The Fungi (G. C. Ainsworth, F. K. Sparrow & A. S. Sussman. 



Eds.) 4A : 5 1 3-582. New York & London. 

— 1975. Coelomycetes. V. Coryneum. Mycol Pap. 138 : 1-224. 

— 1977. Coelomycetes. VI. Nomenclature of generic names proposed for Coelomycetes. Mycol. Pap. 
141 : 1-253. 

— 1980. The Coelomycetes. Kew. 



Swinscow, T. D. V. & Krog, H. 1976. The Usnea articulata aggregate in East Africa. Norw. J. 

Bot. 23:261-268. 
Tibell, L. 1975. The Caliciales of boreal North America. Symb. bot. upsal. 21(2) : 1-128. 

1976. The genus Thelomma. Bot. Notiser 129 : 221-249. 

1978. The genus Microcalicium. Bot. Notiser 131 : 229-246. 

Tobler, F. 1913. Verrucaster lichenicola nov. gen., nov. spec. Abh. naturw. Ver. Bremen 21 : 383-384. 
Tulasne, L.-R. 1852. Memoire pour servir a Fhistoire organographique et physiologique des 

lichens. Ann. Sci. nat. (Bot.) Ill, 17 : 1-225. 
Vainio, E. |Aj. 1887. Monographia Cladoniarum universalis. Pars prima. Acta Soc. Fauna Fl. 

fenn.4: 1-509. 
Vezda, A. 1963. Ph'spdvek k poznani lichenikolnich hub v Ceskoslovensku I. Ceskd mykol. 

17:149-159. 

1970. Pfispevek k poznani lichenikolnich hub v Ceskoslovensku III. Ceskd mykol. 24 : 220-229. 

Viegas, A. P. 1961. Indice de Fungos da America do Sul. Campinas. 

Vobis, G. 1977. Studies on the germination of lichen conidia. Lichenologist 9 : 131-136. 

1980. Bau und Entwicklung der Flechten-Pycnidien und ihrer Conidien. Bibltheca lich., 

Lehre 14 : i-v, 1-141. 

& Hawksworth, D. L. 1981. Conidial lichen-forming fungi. In The Biology of Conidial 

Fungi (G. T. Cole & W. B. Kendrick, Eds.) 1 : 245-273. New York, London & San Francisco. 

Vouaux, L. 1912-14. Synopsis des champignons parasites des lichens. Bull, trimest. Soc. mycol. Fr. 

28 : 1 77-256; 29 : 33-128, 399^194; 30 : 135-198, 281-329. 
Werner, R.-G. 1928. Etude biologique et physiologique du Celidium stictarum (de Not.) Tul. Bull. 

trimest. Soc. mycol. Fr. 44 : 194-205. 

1969. Lichenes et fungi maroccani. III. Etude substratique. Bull, trimest. Soc. mycol. Fr. 

85: 195-204. 

1970. Lichenologie du Prerif meridional et de Fancien detroit Sud-Rifain environment. 

Bull. Acad. Soc. lorr. Sci. 9 : 484-505. 

1972. Lichenes et champignons de la plaine Marocaine. Bull. Acad. Soc. lorr. Sci. 11 : 83-97. 

& Deschatres, R. 1974. Contribution a l'etude des lichens de la Corse. III. Bull. Soc. bot. 



Fr 121 : 299-318. 
Zopf, F. W. 1896. UebersichtderaufFlechtenschmarotzenden Pilze. Hedwigia 35 : 312-366. 
1898. Untersuchungen iiber die durch parasitische Pilze hervorgerufenen Krankheiten 

der Flechten (Fortsetzung). Nova Acta Acad. Caesar. Leop. Carol. 70 : 243-288. 



Host index 

Names of lichen-forming fungi mentioned in the introductory sections and elsewhere are also included 
here. Accepted names are in roman and synonyms in italic. 



Anaptychia ciliaris 37 Bryoria capillaris 79 

Anisomeridium biforme 80 fuscescens 37 

Buellia disciformis 29-30 
punctata 8 1 
Bacidia63 
inundata 81 

rubella 88 Calicium9,80 

s pod odes 81 glaucellum 9-10, 44 

Baeomyces rufus 56 trabinellum 8-10 

Bilimbia spododes 85 viride 42, 44 

Blarneya hibernica 63 Caloplaca caesiorufa 38 



LICHENICOLOUS COELOMYCETES 



93 



cerina38,50-51,66 
ferruginea 38 
flavovirescens 38 
holocarpa 66 
lactea 38 
regalis 84 
vitellinula74 
Cetraria nivalis 72 

sepincola37 
Cetrelia olivetorum 37 
Chaenotheca 44 
brunneola 44 
chrysocephala 44 
subroscida42,44 
trichialis 44 
Chrysothrix chrysophthalma 45-46 
Ciadonia 3, 15-17, 19, 26, 36-37, 39, 41, 87 
acuminata 4 1 
alcicornis 39 
amaurocraea 11,13 
anomaea 1 7 
arbuscula 36-37, 41 
bacillaris 88 
bacilliformis 19-20 
bellidiflora 13 
caespiticia 15, 18-19 
cariosa37 
carneola 17 
cenotea 36 
chlorophaea 17, 19 
ciliata var. tenuis 41 
coniocraea 17,21, 35,41 

conoidea 19 

cornuta 18,41 

cyanipes 1 8 

deformis 1 8 

digitata35 

fimbriatal8,36,39,41 

foliacea39,41 

furcata 25-26 

glauca 18-19 

gracilis 41 

grayi 18 

impexa 4 1 

incrassata 36 

major 1 8 

merochlorophaea 18-19 

norrlinii4\ 

ochrochlora 13, 18 

pleurota 18 

pocillum 36 

portentosa 12,41 

pyxidata 18-19,21,36,41 

rangiferina 82 
f. moribunda 37 

rangiformis 82 

sihatica 1 3 

stellaris 13 

stellata f. adusta 1 2 



subulata 18 
surrecta 13 
sylvatica 4 1 
terrae-novae 13-14 
turgida 13, 18 
uncialis 1 1-14 
f. biuncialis 12 
f. leprosa 1 2 
var. leprosa 1 3 
Cliostomum griffithii 3 
Collema tenax 84 



Diploicia canescens 86 
Diploschistes scruposus 38 
Dirina ceratoniae 86—87 



Evernia prunastri 35-36, 38 



Gyalectidium 75 
filicinum 75 



Haematomma elatinum 77-78 
Heterodea muelleri 35 
Heterodermia speciosa 74 
Hypogymnia bitteriana 35 

encausta 76, 79 

intestiiniformis 16-11 , 79 

physodes 35-37, 83 



Lasioloma 63 

arachnoideum 64 
Lecanactis abietina 79 

subabietina3, 79 
Lecania 80 

fuscella 76 

syringe a 76 
Lecanora36,65,72,84-85 

admontensis 36 

alphoplaca 49 

atra 72 

carpinea 36 

chlarona35, 85 

chlarotera36,65-67,85 

confusa 38 

conizaeoides 35-36 

dispersa38,66,81-82,84 

effusa 84 

expallens38 

glaucellal5,$\ 

laevis 67 

pacifica 37 

pallida 36, 38, 82 

piniperda 75, 81 

polytropa36,38,82 



94 



D. L. HAWKSWORTH 



rubina 14 

rupicola 72 

saligna38,80-81,84 

subfuscata 36 

superfluens 36 

umbrina 81-82 

varia 38 
Lecidea 38 

canescens 86 

enteroleuca 38 

leucocephala C. patelliformis 79 

luteal a 88 

parasema 85 

verruca 3 
Lecidella elaeochroma 38, 80, 85 
Letharia vulpina 37, 73 
Lobaria pulmonaria 4, 14-15, 73 
Lopadium 59,62-63 

flammeum 59 

phyllogenum 75 

tayabasense 59 



Mazosia 86 

melanophthalma 78, 85-87 

phyllosema 86 
Melanelia 84 
Micarea chrysophthalma 45 

nitschkeana38,81,85 



Ochrolechia parella 74 
Opegrapha78,82 

air a var. parella 38 

einerea 85 

lithyrga 82 

niveoatra28-29,78,81 

subsiderella 78 

vulgata 85 



Parmelia50,70,72,83-84,87 
acetabulum 83 
afrorevoluta 35 
borreri4, 14, 36 

caperata 5, 35-36, 5 1 , 56, 67, 7 1 , 76, 80 
conspersa 37 
crinita 35 
exasperata 37 
galbina 35-36 
glabratula37,56,72 

subsp. fuliginosa4, 14-15 
jubala S. cana 79 
laevigata 35-36, 5 1-52, 66 
laevior 82 

mollis y. divaricata 79 
olivacea 37, 83 
pastillifera 36 



perforata 35, 83 

perlata 5, 35-36, 51-52,67,71,83 

pulla37 

pulverulenta 82 

reticulata 51-52 

revoluta 35 

rudecta 37, 76 

saxatilis 35-37, 67, 70, 73-74 

sulcata 4, 14-15,36,51-52,76 

verruculifera 37 
Parmeliopsis ambigua 35 
Peltigera55,72-74,76 

canina 24-25, 54, 76 

erumpens 3 1 

horizontalis 54 

malacea 50, 54 

polydactyla31, 54 

pusilla80 

rufescens 22, 24-25, 54 

spuria 30-31, 33 
Pertusaria29,38 

albescens 38 

hymenea 35 

leioplaca29,38,66 

pertusa29, 35 

puslulata f. glabrata 38 
Phaeophyscia orbicularis 8 1 
Physcia 82 

aipolia36-37,50,56 

stellaris 37, 82 
Physconia 82 

pulverulacea 37, 50, 56-57, 82 
Platismatia glauca 4, 14, 54, 67, 69-70, 72 
Polycauliona regalis 84 
Psorotheciopsis 75 



Ramalina67 

baltica37 

calicaris 37, 69 

dilacerata 69 

fastigiata 69 

fraxinea37,67,69,73 

implectens 75 

siliquosa 37 

sinensis 69 

subgeniculata 37 

yemensis35 
Rhizoplaca chrysoleuca 4, 14-15, 36, 72 
Rinodina exigua 82 

septentrionalis 38 



Solorina crocea 78 
Sporopodium 59 
Squamarina lentigera 36 



Tapellaria 62 



Teloschistes 84 
Thelomma siliceum 48 



LICHENICOLOUS COELOMYCETES 

Verrucaria 82 



95 



unidentified corticolous 8, 59, 73 

foliicolous66 

saxicolous 47^48 
Usnea37,50, 54, 59 

barbata vav.jlorida 54 

filipendula37, 54 

florida35,37,54 



Xanthoria72,77 
fallax77 

parietina37,72-73,81-82 
polycarpa 37 



Zahlbrucknerella africana 73 



Fungus index 

In addition to synonyms cited in this revision, this index contains some synonyms of accepted 
species for which further information is provided in papers cited in the text. New taxa, and the 
page numbers of the principal references to accepted species, are printed in bold, accepted 
names in roman, and synonyms in italic type. An asterisk (*) indicates pages on which there is a 
figure. 



Abrothallus67, 70-71, 87 

bertianus 70, 72 

cetrariae 72 

glabratulae 72 

microspermus 70, 76 

moorei 13 

parmeliarum 70-72, 76 

peyritischii 72 

smithii 70, 72 

suecicus 69 

usneae 58, 72 
Acleistomyces rionegrensis 59 
Alysia 64 

pithospora 66 
Aposphaeria cladoniae 39,41 
var.floerkeanae 39,41 

ramalinae 75 

stenospora 20-2 1 
Arthonia clemens 72 

destruens 72 

epiphyscia 72,81 

fuscopurpurea 72 

glaucomaria 72 

varia 72 
Ascochyta 7-8 

lichenoides 5-6,8 

pisi 7 
Ascochytula lecanorae- Lichenodiplis 

lecanorae 
Asterophoma 7, 8-10, 45 

mazaediicola 8, 9* 



Bachmanniomyces 6, 10-14, 16 

uncialicola3,10, 11-12*, 13-14, 16,87,89 



Campylidium 59 

Celidium fuscopurpureum 72 

insidens 72 

stictarum 73 
Chaetothyriolum puiggarii 75 
Chlorocyphella 59 

aeruginascens 60 
var. convoluta 60 
var. cystidiifera 75 
var. staurospora 63 

foliicola 63 

lichenicola 60 

sub tropica 60 
Clathridium corticola 88 
Conida clemens 72 
Coniosporium pertusariicola = Laeviomyces 

pertusariicola 
Coniothyrium cladoniae 87 

epiphyllum 75 

harmandii 75 

imbricariae - Lichenoconium usneae 

jaapii - Lichenoconium usneae 

lecanoracearum s. str. = Lichenoconium 
usneae 

lecanoracearum auct. = Lichenoconium 
lecanorae 

lichenicola - Lichenoconium lichenicola 
var. buelliae = Laeviomyces pertusariicola 

pyxidatae = Lichenoconium pyxidatae 

ramalinae - Lichenoconium cargillianum 

usneae - Lichenoconium usneae 
Coniscosolen mirabilis 75 
Cornutispora 6, 14-15 

lichenicola 4-5, 14, 15*, 51 

limaciformis 14 



96 



D. L. HAWKSWORTH 



Cristidium pallidum 75 
Cyphella aeruginascens 60 

foliicola 63 

subcyanea 60 



Dacampiosphaeria rivana 73, 78 
Dendrophoma 39 

alcicornaria 39 

lecanorae 75 

p ode ti kola 39 
Dichaena pertusariae 38 
Dichosporium glomerata 73 
Didymella parvispora 73 
Dinemasporium 39 
Diplodia lecanorae = Lichenodiplis lecanorae 

parmeliae 76 
Diplodinaclaudelii 16, 19 

lecanorae = Lichenodiplis lecanorae 

lichenoides = Ascochyta lichenoides 

parmeliae 76 

peltigerae 19, 74, 76 

sands tedei 15-16, 19 

solorinaria 78 

vouauxii ? = Lichenodiplis lecanorae 
Discosporium 1 1 

populeum 1 1 
Dothiorella erumpens 87 



Endococcus 83 

Endophragmiella hughesii 15 

Epicladonia 6-7, 15-22, 89 
sandstedei3, 15,16, 17-18*, 19-20,89 
simplex 16, 19, 20-21*, 89 
stenospora 16, 17*, 19,20,21,22* 

Epicoccum parmeliarum 70, 76 



Griphosphaeria corticola £ 
Guignardia olivieri 73, 8 1 



Hansfordiellopsis lichenicola 75 
Hendersonia lichenicola 87-88 
Herpotrichia 84 
Homostegia lichenum 73 
piggotii 73 



Karsteniomyces 6, 22-25 
peltigerae22,23-24*,31,74 

Keissleriomyces 6, 25-26 
sandstedeanus 25, 26-27* 

Keratosphaera batistae 86 



Laeviomyces 5-6, 26-30, 33 i 
opegraphae 28*, 29*, 78 
pertusariicola 27-28, 29* 



Lecidea cladoniaria 1 3 

irregularis 60 
Leptodothiorella 73 
Leptosphaeria 49 
Lethariicola sipei 73 
Levieuxia borealis 1,33 
Libertella 30 
Libertiella5,7,30-33 

malmedyensis 30, 3 1-32*, 83 

obscurior 76 

peltigerae 30-32 

xanthoriae 77 
Lichenoconium 5-6, 27, 33-37 

boreale 33 

cladon iae%l 

cargillianum 33, 34*, 35 

echinosporum 34*, 35 

erodens3,33,34*,35,36,51 

imbricariae - Lichenoconium usneae 

jaapii = Lichenoconium usneae 

lecanorae 14-1 5, 34*, 35, 36 

lecanoracearum auct. = Lichenoconium 
lecanorae 

lecanoracearum s. str. = Lichenoconium 
usneae 

lichenicola 33, 34*, 36 
var. buelliae- Laeviomyces pertusariicola 

parasiticum 36 = Lichenoconium lecanorae 

pertusariicola 29, 33 

pyxidatae33,34*,36, 37 

usneae 14-1 5, 34*, 35, 37, 58, 87 

xanthoriae 33, 34*, 37 
Lichenodiplis 5-6, 27, 37-38 

lecanorae 29, 37, 38, 66, 74 

lichenicola 37,38 
Lichenomyces lichenum 73 
Lichenophoma 77 

haematommatis 77* 

opegraphae 78 
Lichenosticta 5-6, 38^1 

alcicornaria 39, 40*, 41 

podetiicola 39,41 



Melophia woodsiana 60 
Merismatium lecanorae 74 
Microcalicium 7, 41-44, 7 1 

conversum 42, 43*, 44 

disseminatum 42 

subpedicellatum 10, 42, 43*, 44 
Microdiplodia effusae = Lichenodiplis lecanorae 

ferrugineae = Lichenodiplis lecanorae 

lecanorae = Lichenodiplis lecanorae 

lichenicola! = Lichenodiplis lichenicola 
Microthelia alcicornaria 39, 41 

cargillianum = Lichenoconium cargillianum 

solorinaria 78 
Minutophoma 7-8, 44-46 

chrysophthalmae 44, 45*, 46 



LICHENICOLOUS COELOMYCETES 



97 



Muellerella lichenicola 29, 74 



Nectria heterospora 76 

parmeliae 76 

tenacis 84 
Nesolechia cladoniaria 1 3 

oxyspora 74 
Nicholsoniella 30 

malmedyensis 30 
Nigropuncta 46^9 

rugulosa 3, 6, 46, 47-48*, 49 



Orthidium 59 



Phacopsis clemens 72 

huuskonenii 79 

varia 72 
Phaeoantenariella 78 

lichenicola 78 
Pharcidia dispersa 74 
Phoma 5, 7-8, 1 1 , 44, 49-57, 69, 80, 83 

abietinae 79 

alectoriae 79 

arachnoidea 79 

biformis 80 

caloplacae 50*, 5 1 

caperatae 53, 57, 80 

cladoniae 39 

curvispora 80 

cytospora50,51,52*, 53,55 

dubia 50, 53*, 54 

epiparmelia 80 

epiphyscia 81 

fusispora 8 1 

glaucella81 

herbarum 49 

lecanorae 8 1 

lichenis 82 
f. immersa 82 

parmeliarum 76 

peltigerae 50, 54, 55*, 81 

physciicola 50, 56*, 57 
var. caperatae 80 

ramalinae 67 

truncata = Vouauxiomyces truncatus 

uncialicola 1 1 

usneae 57, 79 

verrucariae 82 

versoniana 83 
Phyllosticta 50, 55 

cytospora 51, 83 

galligena 83 

lichenicola 53, 83 

peltigerae 54-55 

uncialicola 10-1 1, 13 
Phyllostictina 50 



Phymatopsis dubia 54 
Pleospora 49 

hookeri 78 
Pleurosticta 83-84 

lichenicola 83 
Polycoccum 83 

rugulosaria 84 
Psammina 7 

Pseudodiplodia lichenis 84 
Pseudoseptoria 7, 57-59 

donacicola 57 

donacis 57 

usneae 5, 16,57,58* 
Pyrenochaeta 84 

collematis 84 
Pyrenotrichum 7, 59-64, 70, 75 

atrocyaneum 59 

bicolor 59 

filiferum59,63 

foliicola59,63 

irregulare 59 

mirum 59 

podosphaera 59 

splitgerberi 3, 59, 60, 61-62*, 63, 75 

staurosporum 59, 60, 63, 64* 



Rhabdospora antarctica 84-85 

lecanorae 85 

lesdainii 85 

thallicola 85 
Rhagadostoma lichenicola 78 



Schecutia 30 

malmedyensis 30 
Sclerococcum 27, 46 
Scutula epiblastematica 74, 76 

wallrothii 74 
Seimatosporium 88 

lichenicola 87-88 
Selenophoma 57 
Sirococcus 1 1 

lichenicola 1 1 

strobilinus 1 1 
Sirothecium lichenicola = Vouauxiella 
lichenicola 
f. bispora = Lichenodiplis lecanorae 
f. cerinae = Lichenodiplis lecanorae 
Sphaeromma mazosiae 85-87 
Sphaeronema lichenophilum 86-87 
Sphaeropsis cladoniae 87 

scripta 87 
Spilomium 27 

epicladonia 14, 87 

pertusariicola 27, 29 
Sporhaplus rondoniensis 86-87 
SporocadusSl 

lichenicola 87 



98 



D. L. HAWKSWORTH 



rosicola 88 
Stagonopsis22, 84 

lichen is 84 

pallida 22 

peltigerae 22, 55 
Stagonospora25 

paludosa25 

sandstedeana 25 
Stigmidium dispersum 74 



Taeniolella 66 

Telimena foreaui 74 

Tichothecium erraticum 74 

Torula lichenicold = Vouauxiella lichenicola 

verrucosa = Vouauxiella verrucosa 
Trichosperma aeruginosa 60 

cyphelloidea 60 



Verrucaster 16 

lichenicola 88* 
Vouauxiella 3, 6, 64-67 

lichenicola 65, 66-67 

pithospora65,66 

uniseptata 5, 46, 65, 66 

verrucosa 65,67 
Vouauxiomyces 7, 67-71, 72 

ramalinae67,68*,69,75 

santessonii 5, 67, 68*, 69, 7 1-72, 76 

truncatus 5,67, 68*, 69, 70, 7 1 *, 76, 80 



Xeroconium 33 
boreale 33 



Zythia peltigerae 30 



Manuscript received 25 September 1980 



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Titles to be published in Volume 9 



The lichenicolous Coelomycetes. By D. L. Hawksworth 

The genus Callithamnion (Rhodophyta: Ceramiaceae) in the 
British Isles. By P. S. Dixon and J. H. Price 

Parmelia subgenus Amphigymnia (lichens) in East Africa. 

By Hildur Krog and T. D. V. Swinscow 

The genus Selaginella in tropical South America. 

By A. H. G. Alston, A. C. Jermy and J. M. Rankin 



Printed by Henry Ling Ltd, Dorchester 



1 4 MAY 1982 

Bulletin of the 

British Museum (Natural History) 



The genus Callithamnion (Rhodophyta: 
Ceramiaceae) in the British Isles 

Peter S. Dixon & James H. Price 



Botany series Vol 9 No 2 26 November 1981 



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World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) 



Trustees of the British Museum (Natural History), 1981 



The Botany Series is edited in the Museum's Department of Botany 
Keeper of Botany: Mr J. F. M. Cannon 

Editor of Bulletin : Mr P. W. James 

Assistant Editor: Mr J. R. Laundon 



ISSN 0068-2292 Botany series 

Vol 9 No 2 pp 99-141 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 26 November 1981 




M&ftARY 



* GENERAL * 

The genus Callithamnion (Rhodophyta: ( f 4MAY m2 

Ceramiaceae) in the British Isles 

Peter S. Dixon 

Department of Ecology and Evolutionary Biology, University of California, Irvine, 
California 927 17, U.S.A. 

James H. Price 

Department of Botany, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Contents 

Synopsis 100 

Introduction 100 

Aspects of structure and reproduction 100 

Analysis of character validity: adult material 101 

1 . Main axis characters 101 

(a) Integrity of a main axis throughout the thallus 102 

(b) Degree of cortication of the main axis and major lateral axes . . . 102 

(c) Development of lateral filaments from the cortication .... 104 

(d) Cell dimensions 104 

2. Characteristics of major laterals 105 

(a) Branching patterns of major laterals 105 

(b) Number of orders of laterals distinguishable 105 

(c) Reflexion of major laterals 105 

3. Characteristics of peripheral laterals 105 

(a) Branching patterns of peripheral laterals 105 

(b) Occurrence of 'secund processes' 106 

(c) Overall shape and size 106 

(d) Hyaline hairs 106 

4. Reproductive characteristics 106 

(a) Sporangia 106 

(b) The gametangial phase 107 

(c) The carposporophyte 113 

5 Gross criteria of species and groups . . . 113 

The genus Callithamnion Lyngbye 113 

General key to British species , 115 

Accepted species 116 

1. Callithamnion byssoides Arnott ex Harvey 116 

2. Callithamnion corymbosum (Smith) Lyngbye 118 

3. Callithamnion decompositum J. Agardh 119 

4. Callithamnion granulatum (Ducluzeau) C. Agardh 120 

5. Callithamnion hookeri (Dillwyn)S. F. Gray 122 

6. Callithamnion roseum (Roth) Lyngbye 123 

7. Callithamnion sepositum (Gunnerus) P. Dixon & J. Price .... 124 

8. Callithamnion spongiosum Harvey 126 

9. Callithamnion tetragonum (Withering) S. F. Gray 127 

10. Callithamnion tetricum (Dillwyn)S. F. Gray 128 

Rejected species 129 

1 . Rejections from the British flora 129 

2. Entities of uncertain attribution 130 

3. Rejections from the genus 130 

Typifications 131 

Acknowledgements 138 

References 139 



Bull. Br. Mus. nat. Hist. (Bot.)9 (2): 99-141 Issued 26 November 1981 



1 00 P. S. DIXON & J. H. PRICE 

Synopsis 

The algal genus Callithamnion is represented in the British Isles by 10 species: C. byssoides, C. corym- 
bosum, C. decompositum, C. granulatum, C. hookeri, C. roseum, C. sepositum, C. spongiosum, C. 
tetragonum and C. tetricum. There is one new combination: C. sepositum (Gunnerus) P. Dixon & J. 
Price. 

A critical description is provided for the genus and for each species, together with comments on those 
features of structure and reproduction of systematic importance, detailed analyses of the validity of 
these features for taxonomic distinction, and the range of variation encountered in the British Isles and 
in adjacent regions. A general key to the British species is provided, together with details of the 
typification of all entities relevant to a consideration of the genus in the British Isles. 

Introduction 

The genus Callithamnion presents a supreme example of the taxonomic confusion in 
red algae.^The following are the principal reasons for this: 

1 . A lack of understanding of the effects of the environment on an extremely 'plastic' 
phenotype, capable of variation at many different levels. 

2. The lack of data on the extent of that morphological variation and its relation to 
the limits of subsidiary taxa in different geographical areas. 

3. The misunderstanding, misinterpretation and confusion in the application of names at 
different times, by different workers, in different countries. 

The present paper is the beginning of an attempt to rationalize treatment of this group of 
algae, in terms of morphology, development, variation and systematics. 

Our basic orientation has been towards those entities occurring on or reported from British 
shores. As a result of carrying out typifications and dealing with doubtful reports and 
records, it became necessary to refer to entities from elsewhere, mostly in western Europe but 
also in other parts of the Atlantic and Mediterranean. Since the geographical confines of this 
study were envisaged originally as being from North Cape to the Algarve, with treatment of 
the genus in full to be undertaken only for the British Isles, data on Callithamnion in areas 
outside Great Britain and Ireland have been considered only in so far as they affect the 
situation in the British Isles. Treatment of the genus in those areas outside the British Isles is, 
therefore, not complete. 

Aspects of structure and reproduction 

The thallus in species of Callithamnion is of uniseriate filamentous construction, although 
this may become obscured through the development of 'cortication'. Each uniseriate 
filament results from the activity of a single apical cell which divides transversely or 
obliquely. Once formed, a product cell does not divide again in the plane in which it was 
produced, so that the number of cells in an undamaged filament is a reflection of the number 
of divisions of the apical cell. Cells may be uninucleate or multinucleate when mature and, 
apart from rhizoidal cells, each contains several pigmented chloroplasts. The shape and 
number of chloroplasts are modified by age or the conditions in which the thallus is growing. 
Although a product cell does not divide again in the plane of division in which it was 
formed, it frequently does divide once more in a different plane to form the primordium of a 
lateral branch. Each primordium is clearly of lateral origin, although the final disposition of 
branches may appear dichotomous if evection occurs. 

Each thallus in Callithamnion is thus a complex pattern of filaments. A few filaments are 
of unlimited or unrestricted growth, whereas most are of limited or restricted growth. The 
difference between these two types is of a physiological nature, with few easily-definable 
morphological differences. Interconversions can occur, although for many species there are 
characteristic arrangements of filaments of limited and unlimited growth. In certain species, 
other lateral filaments may form which grow downward, investing axes, to form the 
'cortication'. These 'corticating' filaments are composed of pigmented cells and they arise 



CALLITHAMNIONIN THE BRITISH ISLES 101 

from the basal poles of axial cells or from the lowermost cells of lateral filaments. The latter 
cannot be clearly distinguished from 'corticating' filaments as either can readily give rise to 
the other. Nevertheless, the term 'corticating filament' has utility in indicating function and 
position at that time. The extent of cortication varies considerably, even within and between 
individuals of the same species. 

Following the formation of lateral primordia, cells undergo considerable enlargement so 
that the volume of mature cells may be as much as 50,000 times the original volume. This 
enlargement is almost entirely in the basal region of a cell, so that in the mature state the 
lateral filament is attached at the apical pole. Enlargement is associated with development of 
a large central vacuole and as a result the cytoplasm becomes a peripheral lining within the 
wall. Cell size varies considerably, not only with respect to age but also to environmental 
modifications caused by temperature, light intensity, light quality and photoregime. There 
are characteristic patterns of variation in cell size due to interactions between laterals and the 
axes on which they are borne. In some species of Callithamnion this is shown most clearly in 
cells at the base of lateral filaments. These cells are older than cells further along the lateral 
filaments so that they should be larger, but their smaller size indicates that some retardation 
of enlargement has occurred, possibly through proximity to the axial filament. There may be 
other factors involved, since a similar phenomenon occurs in main axes (cf. Dixon, 1971; 
Price, 1978). 

A thallus is attached to the substrate by multicellular rhizoidal filaments which resemble 
'cortication' but differ in that pigmented chloroplasts are absent. Such filaments also 
occasionally develop from the cortication as adventitious outgrowths (Price, 1978). When 
spores germinate, they divide into an upper and a lower cell, the latter forming only the 
primary rhizoid. Subsequently, additional rhizoids are formed, initially from the lowermost 
cells of the axial filament and then from the lowermost cells of lateral filaments. 

This discussion of the developmental morphology of Callithamnion may at first sight 
appear to differ markedly from the descriptions given in recent years by various French 
workers. These use an elaborate system of nomenclature first proposed by Chadefaud 
(1954; see also 1979). In its original form, this system carried the implication that morpho- 
logical categories were rigidly distinct one from another, although in recent publications by 
L'Hardy-Halos (1970, 1971<2, b, c) a more flexible interpretation has been adopted which we 
consider relates better to the realities of morphogenetic plasticity in Florideophyceae. This is 
not the place for a detailed analysis of the French system, and the reader is referred to the 
publications cited for its details. It seems to us that, with few exceptions, the differences of 
opinion between the system which we have used and the French system are now largely 
semantic. 

In its vegetative morphology, Callithamnion resembles some other genera of the 
Ceramiaceae, from which it differs in reproductive details. Callithamnion and its immediate 
allies, the 'Callithamnion-GruppQ' of Kylin (1956), are characterized by the lateral 
intercalary position of procarps. After fertilization, the support cell divides to produce an 
auxiliary cell. Once contact has been established with the carpogonium, the auxiliary cell 
gives rise to gonimoblast initials. These form the filaments of the carposporophyte; the latter 
may be single and entire or deeply lobed. In either event, each cell of the gonimoblast 
develops into a carposporangium which produces a single carpospore. 

Various other types of spores occur in the genus. Tetrasporangia are probably of the widest 
occurrence. Structures producing more than four spores have been described and these 
appear to be parasporangia (cf. Drew, 1939), while structures producing only two spores 
have been observed. Claims for the occurrence of mature, functional monsporangia cannot 
be substantiated. For comments on polysporangia, see p. 106. 

Analysis of character validity: adult material 
1. Main axis characters 

The characteristics of main axes demand careful examination to ensure that the structure 



02 



P. S. DIXON & J. H. PRICE 



under investigation is the main axis. Major branches may resemble the primary axis but the 
identity should not be assumed without question. Analysis of main axis characteristics 
requires a whole thallus, but herbarium specimens of Callithamnion are not always 
complete. Thalli were often 'pruned' to produce a specimen with a greater aesthetic appeal, 
while thalli occurring on animals are usually stripped from the substrate during either 
collection or preservation and thereby often damaged. 

(a) Integrity of a main axis throughout the thallus 

Many authors have characterized species of Callithamnion by the degree of distinctness of 
the main axis, differentiating between species in which a main axis is easily distinguishable 
and those in which it is not. This feature depends in part on whether branching is obviously 
lateral, and on the extent to which evection occurs so that the branching is of a more 
dichotomous aspect. From personal observations, the degree to which a main axis remains 
identifiable in a species is highly variable and the feature is of little value of taxonomic 
discrimination. 

(b) Degree of cortication of the main axis and major lateral axes 

Virtually all treatments of the genus Callithamnion have made reference to cortication. Only 
rarely has this feature been discussed with any degree of precision or with reference to its 
variability. Appreciation of the relationship between degree of cortication and environ- 
mental factors is a relatively recent development (Boddeke, 1958; Harris, 1962; Price, 1978). 





B 



Fig. 1 A, Callithamnion tetragonum: a well-corticated axis showing a primary lateral and two 
adventitious axes arising from the cortication (x 70). B, C. sepositum: transverse section of a 
well-corticated axis (x 35). C, C. byssoides: origin of cortication by down-growth from the basal 
cell of a primary lateral (x 250). 




CALL1TH AMNION IN THE BRITISH ISLES 



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104 



P. S. DIXON & J. H.PRICE 



These authors comment that cortical and adventitious development is greatest in entities 
tolerating or exposed to wave action. This increase in cortication density in such entities 
appears to result from two causes. There is an increase in the formation of the downward- 
growing filaments which constitute cortication. Also, without exception, entities occurring 
in conditions of strong wave action display cells (in the main axis and major laterals) which 
are short compared with their diameter. As a result, the amount of cortication is increased 
while the surface area to be covered is less than in an entity where cell enlargement is greater 
(Fig. 1). 

(c) Development of lateral filaments from the cortication 

The down-growing corticating filaments can give rise to adventitious lateral filaments and 
the occurrence of these has been used by some authors (e.g. Rosenvinge, 1924) to 
characterize certain entities. Maximum expression of this feature occurs in those entities 
tolerant of or exposed to severe wave action, where cortication development is greatest. The 
feature is of little general use as a strictly taxonomic criterion in the genus Callithamnion 
(Price, 1978). 

(d) Cell dimensions 

The dimensions or length-to-breadth ratios of cells have been used extensively for discrimi- 
nating between entities in the genus Callithamnion (e.g. Kylin, 1907; Newton, 1931; 
Boddeke, 1958) even though some authors (e.g. Rosenvinge, 1924) have questioned the value 
of this feature. It has been demonstrated (Dixon, 1971) that there are elaborate patterns of 
cell enlargement in species of Callithamnion, with regard to both main axes and principal 
laterals (Fig. 2). Patterns of variation in length-to-breadth ratios are even more complex 
(Fig. 3). In Pleonosporium, which is of similar morphology to Callithamnion, variation in 
cell size has been related to a complex interaction of light intensity and duration (Murray & 
Dixon, 1972, 1975). There are indications that cell size variation in Callithamnion is 
controlled by similar environmental factors. 

The degree of variation in cell size makes it impractical except in certain instances to use 
this feature for taxonomic discrimination. A further implication of the available studies is 
that if cell sizes or ratios are quoted for any entity, the positions of the cells measured must be 
precisely cited. For further detailed consideration, see Price (1978). 




APEX 



t 1 r — r 

20 30 40 50 

SEGMENT NUMBER 



60 BASE 



Fig. 3 Callithamnion corymbosum: histogram showing the length/breadth ratio of every cell 

along the principal axis of a small thallus. 



CALLITH AMNION IN THE BRITISH ISLES 1 05 

2. Characteristics of major laterals 

The most important characteristics of the major laterals in the genus Callithamnion, from 
the taxonomic standpoint, are in their branching patterns and arrangement. As previous 
authors (e.g. Boddeke, 1958; Harris, 1962; Price, 1978) have noted, differences in branching 
patterns may occur in the same entity in different reproductive states so that the condition of 
material must always be indicated in discussion of this feature. The arrangement of major 
laterals in Callithamnion is frequently described as 'phyllotaxis', a term not strictly 
applicable to the algae in the absence of leaves. The alternatives are clumsy or even more 
inaccurate so that, despite the reservations on its use, we employ 'phyllotaxis' for the 
arrangement of major laterals. 

(a) Branching patterns of major laterals 

Of the several arrangements of filaments that are related to patterns of branching and have 
been employed in taxonomic discrimination within the genus, two are most significant. 

1. The relationship between two individual filaments derived from a common cell lineage; 
the arrangement may be alternate or pseudodichotomous. 

2. The relationship between successive lateral filaments along an axis; these may lie in a 
single plane (giving a distichous or biseriate arrangement) or be distributed radially 
(giving a polystichous or spiral arrangement). 

Both are potentially useful criteria, although there can be variations in different parts even of 
a single specimen. Some diagrams illustrating 'phyllotaxis' have been published by L'Hardy- 
Halos (1970) and more are being prepared for use elsewhere. 

(b) Number of orders of laterals distinguishable 

With few exceptions (e.g. Callithamnion caudatum), the number of orders of laterals is 
highly variable depending on a complex of factors, such as the species, the age of the thallus, 
its vigour and the environment in which it occurs. This feature is therefore of little 
taxonomic value for specific discrimination. 

(c) Reflexion of major laterals 

The tendency of major laterals, particularly of older basal laterals, to be reflexed has been 
used as a characteristic of certain entities (e.g. Westbrook, 1927). The feature is highly 
variable and of little taxonomic value. 

3. Characteristics of peripheral laterals 

The thalli of species of Callithamnion are richly branched and the overall form is a reflection 
of the branching pattern of the constituent filaments. The form of the thallus is determined 
not only by the arrangement of major laterals, discussed in the preceding section, but also by 
the arrangements which prevail in the ultimate portions of the constituent filaments. It has 
become apparent from our investigations that considerable taxonomic dependence can be 
placed on the morphology, branching pattern and spatial arrangements of what we term the 
peripheral laterals, so-called because they contribute to the periphery of the thallus. 
Previous workers with the genus have made extensive use of this feature but, in almost every 
case, each has used a different term by which to refer to it. A cursory survey of the literature 
disclosed more than 50 terms which had been applied, of which the most widely used were 
probably ramuli, ramelli, branchlets, pinnae, and pinnules. The use of different terms, often 
with variant applications, has produced an impossibly complex terminological situation and 
it is beyond the scope of the present paper to discuss every usage. 

(a) Branching patterns of peripheral laterals 

The branching patterns of major axes can differ markedly from those of peripheral laterals, 



106 P. S. DIXON & J. H. PRICE 

such differences often being associated with the formation of reproductive structures. Never- 
theless, branching patterns at the peripheral level are usually sufficiently distinct and 
characteristic for taxonomic discrimination. 

(b) Occurrence of 'secund processes' 

One particular aspect of the branching pattern of peripheral laterals relates to the occurrence 
of ultimate lateral filaments which consist only of a single cell. These have been termed 
'secund processes' (e.g. Newton, 1931) and used for taxonomic discrimination. Our investi- 
gations show that these structures are nothing more than early developmental stages of other 
organs or laterals. 

(c) Overall shape and size 

The overall shapes and sizes of peripheral laterals and of their constituent cells have been 
used by many authors (e.g. Rosenvinge, 1924; Newton, 1931; Feldmann-Mazoyer, 1941) in 
diagnoses and for discrimination. In our experience, the sizes of entire peripheral laterals are 
very variable although the shapes and relative sizes of constituent cells are more constant. In 
addition, other features of a peripheral lateral, such as the number of cells which it contains 
and its pattern of branching, are particularly significant. 

{d) Hyaline hairs 

Apical or sub-apical hyaline hairs occur frequently on peripheral laterals in some species of 
the genus. The function of these structures is unknown and, although widespread, their 
occurrence and abundance are subject to great variation. A further difficulty preventing the 
use of these structures for taxonomic discrimination is that they may be damaged or 
destroyed by fixation or the preparation of herbarium specimens. For further detailed 
considerations, see Price (1978). Culture conditions may induce or permit the formation of 
such hairs where field conditions would not. Rosenvinge (1924) found hairs so rarely in 
Callithamnion byssoides (as C furcellariae) that he suggested so distinguishing the taxon 
from C corymbosum; Rueness & Rueness (1980), by contrast, frequently observed 
terminal hyaline hairs on female gametophytes of C. byssoides in culture. 

4. Reproductive characteristics 

Reproductive characteristics tend to be relatively stable but the structures are ephemeral and 
their use as a source of characteristics has been kept to the minimum in this general 
treatment. 

(a) Sporangia 

Monosporangia have been reported in various species of Callithamnion. Field reports are of 
immature tetrasporangia, while reports in culture must be dismissed as artifacts so that we 
have ignored monosporangia for taxonomic purposes. 

Bisporangia are produced inconsistently, even in entities where they have been reported, 
sometimes alone or mixed with tetrasporangia. Their occurrence cannot be relied upon as an 
absolute criterion. For their significance, see Guiry (1978). 

Tetrasporangia are very common and their position, size and shape have been much 
employed taxonomically in the genus Callithamnion, although these characteristics are 
minimally useful. 

A few instances of sporangia containing more than four spores have been detected in 
British species of Callithamnion. These structures can be regarded as parasporangia rather 
than polysporangia (sensu Drew, 1939). The structures are of rare occurrence and the 
position is confused because C decompositum (which here seems habitually to produce 
parasporangia as the consistent reproductive structure) has previously been overlooked in 



CALLITH AMNION W THE BRITISH ISLES 



107 



the British Isles (Price & Tittley, 1978). Until it can be confirmed that British C. hookeri does 
produce parasporangia, as in Scandinavia, or does not, their use as a taxonomic distinction is 
best avoided. 

(b) The gametangial phase 

Previous authors have made considerable use of structures of the gametangial phase in 
distinguishing between species. The principal difficulty is that these structures are com- 
paratively ephemeral. For this reason, no attempt has been made to utilize data from the 
male and female structures or carposporophyte in the present general key. Keys for morpho- 
logical states other than the vegetative and for ecological characteristics are certainly useful. 
Keys involving distinctions between the reproductive states of all accepted species will be 
presented elsewhere (Price, in preparation) although available descriptions are offered in 
species characterization. As otherwise indicated, periodicity of reproductive structures is 
less useful and is discussed but not employed. 

In the male organs, group distinctions based on the degree of branching in the sperma- 
tangiophore system and the positions of spermatangia are reasonably reliable. Positions of 
the spermatangial clusters and their mature form have also been employed. This is not a new 
approach; many authors have attempted to utilize these features. 






ABC 

Fig. 4 Cell patterns in the carpogonial branches of different species of Callithamnion. A, C. 
corymbosum, B, C. byssoides. C, Material identified as k C. tripinnaturrC . After Miranda (1934). 
Diagrammatic, not scaled. 

The employment of cell spatial arrangements and alignment in the carpogonial branch 
owes much to observations by Miranda (1934), but has been subsequently developed by 
other authors. Apart from the ephemeral nature of the material, difficulties of accurate 
observation of the arrangement of cells precludes use of the data at a principal distinguishing 
level, but we have employed these data in other contributory capacities. The original 
concepts put forward by Miranda are shown diagrammatically in Fig. 4; our suggested 
modifications of or additions to these, with terminological clarification, appear as Fig. 5. 
Since there is some variation in arrangement and cell relationship, it is highly likely that 
these aspects of the carpogonial branch depend at least on the relative position of the branch 
on the thallus, on the bearing support cell, and on the bearing axis. The general position 
away from functioning apices may reflect the level at which vegetative apical dominance is 
no longer inhibitory. 

Sizes and shapes of carpogonia, trichogynes, and other cells of the procarp all appear too 
variable to be of taxonomic value at the species level. Trichogyne ramification, not often 
reported, has been noted by us in a few entities, but its inconsistency suggests aberrations 
and/or environmental influences. It is possible that different patterns of cell size relationship 
would emerge for different species from a statistical survey of the carpogonial branch, but the 
taxonomic utility of this is doubtful. 



108 P. S. DIXON & J. H. PRICE 

Fig. 5 Suggested modifications and additions to cell spatial arrangements and alignment types 
recognized in carpogonial branches by Miranda (1934): variations especially emphasized. (All 
measurements in urn.) Pp. 109-1 12. 

5A, Callithamnion tetragonum, straight form usual to species: Carpogonium width (c/w) 1 5. 

Trichogyne length (t/1) 195. 

5B, C. tetragonum, intermediate form near zigzag; c/w 7; t/1 99. 

5C, C. tetragonum, intermediate form near curved: (i) c/w 1 1 ; t/1 48; (ii) c/w 9; t/1 64. 

5D, C granulatum, zigzag form usual to species: (i) c/w 18; t/1 239; (ii) c/w 1 8; t/1 7 1 . 

5E, C.sepositum, curved form usual to species: (i) c/w 18; t/1 75;(ii)c/w 17; t/1-; (iii) c/w 26; t/1 106. 

5F, C tetragonum, straight form with extruded 2: c/w 14; 1/1*171. 

Variations perceived probably depend on relative positions of branch on the axis, on the support cell, 
and on the overall thallus. They are therefore a reflection of the balance between constraints and 
overall space available. Types recognized certainly represent arbitrary divisions of a continuum of 
variation. All figures presented are taken from actual carpogonial branches; dimensions given for the 
carpogonial width and trichogyne length. Arrows and curves = space relations of cells determined by 
focal-planing. Symbols: R = bearing axis cells or line of them; S = support cells (S,; S 2 ); 1 = cell 1 of 
carpogonial branch; 2 = cell 2 of c/b; 3 = cell 3 of c/b; 4 = carpogonium with trichogyne. 

Straight form 
(Callithamnion corymbosum-type of Miranda, 1934; 'straight'-type of Harris, 1959, 1962, 1966). 

Orientation of cells 1 ^2 usually at right angles to long axis of bearing axial cell (walls of 1/2 

therefore parallel to that axis), as in Fig. 5A (example) and Fig. 4A (diagrammatic). Cells 1 »2 

and 2 -►3 are in linearly parallel arrangement, carpogonium (4) being variously arranged on 3 

(Fig. 5 A). Form occurs usually in C. tetricum, C. sepositum, C. corymbosum, C roseum, and C. 

tetragonum (Fig. 5A), sometimes in C. granulatum, C. hookeri, C. byssoides and C. spongiosum. 

Zigzagform 

(Callithamnion byssoides-type of Miranda, 1934; "zigzag'-type of Harris, 1959, 1962, 1966). 

Compound curve or zigzag (Fig. 4B, diagrammatic). Orientation 1 +»2 inclined upwards or at 

outward angle to long axis of bearing axial cell (Fig. 5D). Form occurs usually in C. byssoides, and C. 
granulatum (Fig. 5D), sometimes in C. hookeri, C spongiosum, C roseum, C tetricum, C 
sepositum, C tetragonum and C. corymbosum. 

Curved form 

(Callithamnion \tripinnatum? '-type of Miranda, 1934) 

Single curve ('U'-shaped), often in more than one plane by lateral curvature (Figs 4C/5E). Form 

occurs usually (?) in ?C tripinnatum, often in C. sepositum (Fig. 5E), and probably sometimes in all 

known species. 

Variations of 'standard' forms 

(i). Parallel status of 1 ^2/2 ^3 depends on position of pit connexion between cells 2 and 3. 

If the connexion is high on 3 and 4 (carpogonium) is large or in cramped position, depression or offset 
position of 3 will cause the relationship 

4 

^^3 

to occur (as in Fig. 5B, where 4 (carpogonium) rises towards viewer out of the paper, but would 

obscure detail if drawn fully so). This is very close to the form attributed by Miranda (1934) to C 

byssoides-iype (Fig. 5D; here in examples of C. granulatum). 

(ii). If cell 2 is depressed from the 'normal' C. corymbosum-typc position, and pit connexions 1/2 and 

2/3 are high on 2, then the 'U'-shape of Miranda's (1934) 'tripinnatum' -type is approached, as is 

usual in C. sepositum (Fig. 5E) and occasional in C. tetragonum (Fig. 5C). 

(iii). Arrangement of the carpogonial branch is not always planar, as is clear from the above and from 

curvature diagrams accompanying parts of Fig. 5. Angling to the vertical or into more than one plane 

is common to usual, and can confuse the pattern as, e.g. in Fig. 5F. There, a C tetragonum branch 

with 1 +~2 ► 3 all still parallel has 2 extruded by pressure to jut out from the usual line. 

Otherwise, the branch is clearly of C. corymbosum-type (straight), the periphery of the lateral branch 
system bearing the carpogonial branch being basal to the figure. Harris's (1959, his text-figure 13C) 
intermediate type may have been due to this non-planar situation. 



CALLITHAMNION IN THE BRITISH ISLES 



109 




5B 





110 



P. S. DIXON & J. H. PRICE 




Support cell 
not visible 




Fig. 5 Suggested modifications and additions to cell spatial arrangements and alignment types 
recognized in carpogonial branches by Miranda (1934): variations especially emphasized. (All 
measurements in urn.) Full explanation on p. 108. 



CALLITHAMNIONIN THE BRITISH ISLES 



111 




2/ 4 

If flattened 



probably 



Si 4 



5Eii 



Trichogyne 
fractured in 
position shown 



\ s 




o: 



Line of 

peripheral 

lateral 



112 



P. S. DIXON & J. H. PRICE 



5F 




Fig. 5 Suggested modifications and additions to cell spatial arrangements and alignment types 
recognized in carpogonial branches by Miranda (1934): variations especially emphasized. (All 
measurements in urn.) Full explanation on p. 108. 



CALLITHAMNION IN THE BRITISH ISLES 1 1 3 

(c) The carposporophyte 

Events occurring after fertilization have a standard sequence. It appears that the extent of 
development and number of cells involved in primary and secondary gonimoblast have a 
relative constancy that is useful at a contributory level in species distinction. To some extent, 
basic differences in the early carposporophyte development are reflected in lobing of the 
carposporophyte at early maturity (carposporangial production). However, subsequent 
environmental and morphogenetic effects obscure or override the earlier patterns and 
produce variations in form so that species patterns cannot be distinguished. We have 
employed such criteria with circumspection in the following treatment. Sizes and shapes of 
carposporangia and carpospores are of little taxonomic value. 

Development of a rudimentary vegetative 'involucre' around the carposporophyte occurs 
in some of the more delicate species. It is referred to as necessary in the treatments that 
follow, but it is inadvisable to employ this essentially ephemeral phenomenon as a 
fundamental criterion. 



5. Gross criteria of species and groups 

In addition to particular features used as systematic criteria, numerous other gross criteria 
are cited in diagnoses. Generally, these gross criteria are of little importance and we do not 
employ them for taxonomic distinction. 

Periodicity of upright growth is similar in all species, with one possible exception. It is 
probable that the less robust forms rarely persist as recognizable basal fragments through the 
winter, although it is difficult to be sure of that. Robust, exposed shore forms show rather 
abrupt dieback to variable basal portions in the usual conditions of late autumn or early 
winter. Those thalli which do persist have lost peripheral or even whole branch systems and, 
although still recognizable, have a very different appearance from the luxuriant summer 
growth. One exception, Callithamnion tetragonum, tends to persist as recognizable growths 
on laminae of the Laminariales in some parts of its range. In some years and locations, C. 
tetricum behaves similarly. Although contributory, this behavioural difference is not of 
prime taxonomic importance. 

Statistically, characteristic growth forms can be recognized for most species of 
Callithamnion. Unreliable as primary taxonomic criteria, these are nevertheless very useful. 
The growth forms are a consequence of growth vigour, robustness, texture, frond outline, 
branching pattern, and other unquantifiable phenomena. In some species, external form is 
almost indescribable but still highly characteristic. Species of Callithamnion rarely form 
homogeneous populations with individuals closely-knit enough to possess a recognizable, 
distinct population morphology. Two species form occasional exceptions to this. C. 
roseum populations mainly manifest what is merely a multiplication of the individual 
plant characteristics; C. scopulorum (now C hookeri) has the most recognizable population 
morphology that is rather more than merely the sum of many individuals. Many authors 
have employed the term 'turf for this. The terms is not entirely satisfactory but it is at least 
evocative. In northern parts of the British Isles, exposed shore species (principally C. 
sepositum) sometimes grow so densely as to produce what has been equally inaccurately 
described as 'spongy turf. 



The genus Callithamnion Lyngbye 

Lyngbye, Tent. Hydr. Dan.: 123 (1819). 

Phlebothamnion Kiitzing, Phyc. Gen.: 374 (1843). 

Poecilothamnion Nageli in Neue Denkschr. Allg. schweiz. Ges. ges. Naturw. 9 (un-nod. Art. 

2): 202 (1847). 
Leptothamnion Kiitzing, Sp. Alg.: 896 (1849). 



114 P. S. DIXON & J. H. PRICE 

Dorythamnion Nageli in Sber. bayer. Akad. Wiss. 1861 (2) : 344 (1862). 
Ceratothamnion J. Agardh in Acta Univ. lund. 28 (6) : 35 (1892). 
Aglaothamnion Feldmann-Mazoyer, Rec. Ceram. Med. Occident.: 451 (1941). 

Thallus of variable shape, colour and size; colour rose-pink to muddy yellow-brown, varying 
with species, vigour and habitat in natural conditions, between specimens and within a single 
specimen; size and shape vary with species, specimen and habitat conditions; filamentous, 
axes uniseriate, to 400 mm in length, variably but always copiously branched; branching 
monopodial, although some species are peripherally flabellate from pseudodichotomous 
evection; sometimes corticate by down-growing filaments derived from cells of axis or basal 
cells of primary laterals, less often from subsequent order laterals; cortical development and 
extent variable according to species, specimen and habitat conditions, involving main axis 
only, or all orders of major axes; adventitious branching from cortication in a few species, 
most dense in exposed shore specimens where a densely interwoven continuous thick matrix 
is formed over the cortication; axial widths over cortication reaching 5 mm; normal 
branching alternate, spiral or distichous, or approaching dichotomy in aspect; irregularities 
occur occasionally in all species, with some variation with reproductive state; characteristic 
form of the peripheral laterals occurs in both determinate smaller apical or peripheral 
laterals (commonly referred to as 'ramuli') and indeterminate higher order laterals (in most 
species), where the latter levels represent the current apices of the branching system and 
contribute to the form of the thallus periphery; active main apices (main axes and major 
laterals) either buried within local peripheral lateral development or clearly emerging above 
that development according to species; apical cells of peripheral laterals gradually- to 
abruptly-domed cylinders, of variable length according to species, or cones; apical (or 
subsequently sub-apical) unicellular hyaline hairs irregularly present in active (younger) 
parts of frond in many species, hairs commonly and often suddenly deciduous; cells length- 
to-breadth ratios variable within the range 16 : 1 to 1 : 1 for all parts of the vegetative thallus; 
cells multinucleate with uninucleate immediate products of apical division, or uninucleate 
throughout, according to species; chloroplasts irregular and/or discoid in young cells, 
forming irregular masses or ribbons by division and coalescence when older; chromosome 
numbers commonly variable in the range n = 28-33, with extremes to n = 90-100. 

Gametangial thalli commonly dioecious, occasionally monoecious in many species, 
habitually in one. Spermatangia arise either in separate cushions, formed on cells of 
outermost peripheral laterals, one or more per cell, cushions sometimes coalescing; or in 
secund rows on adaxial surfaces of peripheral laterals, spreading when luxuriant to sides and 
sometimes abaxial surfaces, finally encircling more mature, usually basal, parts; vigorous 
production of spermatangia may lead to additional development from cells of major 
branches of all orders in lateral systems, as in Callithamnion corymbosum or C. hookeri. 

Procarps formed normally just behind or near apices of active axes and major indeter- 
minate laterals, but may occur on lower branches in thalli with more vigorous reproductive 
development; early procarp development with paired support cells, exceptionally with only 
one, formed laterally on bearing axis; carpogonial branch developing on one support cell or 
on both support cells, four-celled, including carpogonium; cell arrangement straight, 
U-shaped, zigzag, or any intermediate of these; carpogonium relatively small, 6*5-26 urn in 
breadth, but with trichogyne of very variable dimensions and shape, curved, ramified, 
nodular, swollen, uneven-sided, or bulbous-tipped, 47-242 urn in length; trichogyne 
degenerating after fertilization, the carpogonium transferring a derivative of the fertilization 
nucleus by a connecting cell, or directly, to an auxiliary cell formed by division from the 
original support cell; following nuclear transfer the auxiliary cell dividing to produce a 
central cell and a foot cell, central cell developing two, less often three, gonimoblast initials, 
all producing few-celled gonimoblast filaments; one or two lobes of the gonimocarp 
developing on the ultimate cell of the gonimoblast, one only on the penultimate cell of the 
gonimoblast and on all subsequent cells; gonimolobes rounded, conical or uneven, varying 
according to species, environmental factors and state of maturity; carposporophyte very 



CALLITH AMNION IN THE BRITISH ISLES 1 1 5 

rarely sheathed by enveloping vegetative filaments; carposporangia of variable dimensions at 
maturity, commonly 25-70 urn in diameter. 

Tetrasporangia ellipsoid or ovoid, less often subspherical to spherical, variable in size and 
shape to 130 x 120 jim, formed adaxially on cells of peripheral laterals, usually one or two 
per cell; bisporangia usually exclusively on particular plants {Callithamnion byssoides) but 
sometimes intermixed with tetrasporangia in C. byssoides and C. hookeri; parasporangia 
reported elsewhere, not often authenticated in Britain, except in C. decompositum. 



General key to British species 

Price (1978) has correctly indicated that ecology should be the initial approach to a 
utilitarian key aimed at field use with fresh material. Here, with all the complications of 
older herbarium or preserved material and sparse ecological data, this is not practicable. 
Nevertheless in the present key an attempt is made to integrate thallus or population 
characteristics likely to have been noted or obvious from morphological state even in 
individuals out of their normal environment. Use is also made of some data of reasonable 
reliability derived from customary habitat preferences. All these aspects have been utilized 
by past authors but an integrated key which does not place too much dependence on any one 
criterion has not been attempted previously. Keys specific to ecology and to reproductive 
states are being published elsewhere (Price, 1978; Price, in preparation). 

1 Laterals of limited growth not overtopping bearing axis 2 

Laterals of limited growth overtopping the axis on which they arise 3 

2 Cells uninucleate; cortication not commonly (although potentially) giving rise to an inter- 

woven matrix of adventitious laterals; thallus almost never coarse, shaggy, or rope-like 

C. hookeri(p. 122) 

Cells largely multinucleate; interwoven matrix of adventitious laterals arising from 
cortication in at least the lower 7/8ths of the thallus; thallus always coarse, shaggy, and 
rope-like in mature specimens, although less so if sublittoral . . C. tetricum (p. 128) 

3 Lateral rudiments arising from the first product of a principal apical cell 4 

Lateral rudiments arising on or after the second product of a principal apical cell ... 8 

4 Apical cells of limited growth peripheral laterals rounded terminally into a dome, sometimes 

strongly tapered 5 

Apical cells of limited growth peripheral laterals very small (12 ± 3 urn long), conical 

C. tetragonum (p. 127) 

5 Cell lengths not exceeding 5 x breadth in any part of the thallus; ramification below lateral 

branch system level strictly pseudodichotomous or not at all so, never mixed ... 6 
Cell lengths exceeding 5 x breadth in older axes and major laterals; ramification below 
lateral branch system level strictly pseudodichotomous . . . C. corymbosum (p. 1 18) 

6 Cells uninucleate; interwoven matrix of adventitious laterals from cortication usually absent 

C. hookeri(p. 122) 

Cell multinucleate; interwoven matrix of adventitious laterals from cortication develops 
over at least most of main axes and principal laterals 

7 Apical clusters of lateral systems > 700 urn across; peripheral laterals (containing more than 

7 cells) with their greatest width well above their base or parallel-sided for much of their 
length; ramification pinnate; common on exposed shores in the sector from south Ireland 

and south Wales northwards round to Yorkshire C. sepositum (p. 124) 

Apical clusters of lateral systems < 700 /im across; peripheral laterals (containing fewer 
than 6 cells) with greatest width at or near their base; ramification pseudodichotomous; 
either rarely recorded anywhere on exposed shores or rare in the north of the British Isles 9 

8 Cells multinucleate; cells near apices > 30 urn diameter . . C. roseum (p. 123) 
Cells uninucleate; cells near apices < 20 um diameter .... C. byssoides (p. 1 16) 

9 Peripheral laterals with more than three cells above last ramification present in most lateral 

systems; common on exposed shores in the southern half of the British Isles 

C. granulatum (p. 120) 

Peripheral laterals with never more than 2 cells, most usually only one cell, per lateral 



116 P. S. DIXON & J. H. PRICE 

above last ramification anywhere in the periphery, detailed peripheral branching pattern 
'aping' C corymbosum; probably widespread on exposed shores but sparse and rarely 
authentically recorded C. spongiosm (p. 126) 

Note: C. decompositum (p. 1 19) is omitted from this key, although described as fully as possible below. 
Comparative information does not permit effective keying with so little material available. The 
presence of parasporangia and strictly distichous apical arrangement indicate that confusion is 
possible only with C. hookeri or Compsothamnion. In the forms of Callithamnion hookeri in which the 
indeterminate apices are not or only just overtopped by determinate laterals, there is commonly 
discernible variation somewhere on the plant from the distichy by which many indeterminate branch 
system apices may resemble those of C. decompositum. The tetrasporangia in C decompositum are a 
mixture of sessile (predominantly) and pedicellate (very occasionally), differing from the consistently 
terminal tetrasporangia on pedicellate laterals in Compsothamnion. The latter is also more delicate and 
less luxuriantly branched than Callithamnion decomposition. 



Accepted species 

1. Callithamnion byssoides Arnott ex Harvey in Hooker, Eng. Fl. Sir J. E. Smith, 
5:342(1833). 

Callithamnion arnottii Trevisan, Nomen. Alg. ... 1 : 77 (1845) 
Callithamnion fur cellar iae J '. Agardh, Sp. Gen. Ord. Alg. 2 (1) : 37 (1851). 
Callithamnion hiemale Kjellman ex Kylin, Stud. Alg. schwed. Westk:. 170 (1907). 

The accepted binomial, Callithamnion byssoides, may by antedated by two binomials. Callithamnion 
arachnoideum C. Agardh, Sp. Alg. 2(1): 181 (1828), was described from the American coast of the 
north Atlantic. Although some American material resembles closely the specimens from the eastern 
shore of the north Atlantic, further detailed study is needed before conspecificity is accepted without 
question. Ceramium tenuissimum Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 132 (1828), was 
described from European material, but exact typification of this binomial is not possible (see p. 1 36). 

Thallus of wide colour range, from pink to brick red; to 70 (-120) mm in height, bushy, 
flaccid, rarely with easily recognizable main axis, although usually only one exists, less often 
several more or less equivalent major axial branches diverge just above attachment; main 
axis to 120(-180)|im in breadth, commonly ecorticate throughout, less often with a few 
basal, down-growing, corticating filaments from lower branches; axial cells narrowly 
elongate, length-to-breadth ratios commonly 5 (or more) to 1 with detected range 300-1 3* 19 
to 1; branching alternate, monopodial throughout, with laterals arising irregularly in 
direction and number; laterals of variable length in lower thallus, of more even length in 
upper; uppermost larger laterals and peripheral laterals sparsely or not branched, long, 
flaccid, flagelliform; insertion appearing distichous in lower regions, but becoming clearly 
spiral towards apices of branching system; peripheral laterals usually unbranched, borne in 
proximal 3/4 of bearing penultimate laterals but forming small corymbose or uneven clusters 
near apices in distal 1/4 of bearing laterals, commonly straight, rarely incurved; apical 
development in lateral systems not pseudodichotomous, but apical cell surpassed by 
peripheral laterals; cells of laterals cylindrical, of similar length on the same lateral; apical 
cells of peripheral laterals elongate domed cylinders, 52*59 ± 17*17 urn in length, 
12*2 ± 3*33 urn in basal breadth and 5*49 ± 2*24 urn in apical breadth, with length-to- 
breadth ratio of (2*5-) 3*5-5*5 to 1; peripheral laterals contain (1-) 2-5 (-7) cells, 51-430 urn 
in length, 7-26 um in greatest breadth, with position of maximum breadth usually above 
base. 

Gametangial thalli dioecious. Spermatangiophores once-branched, sometimes with 
rudiments of second lateral, often several on each cell of peripheral laterals; spermatangio- 
phore branches 4 to 5 cells in length; 1-5 spermatangia on each cell, 6x8 um, but variable in 
size. 

Procarps formed behind apices of major axes and branches, also spreading towards the 
base in very fertile material, with two support cells but sometimes only one; carpogonial 



CALLITH AMNION IN THE BRITISH ISLES 1 1 7 

branches zigzag or straight, on each support cell, but fertilization usually only unilateral; 
two gominoblast initials from each auxiliary cell, first-developed producing two-celled 
primary gonimoblast, second-developed forming one-celled primary gonimoblast, two 
gonimolobes developing from the apical cell of first-developed primary gonimoblast, with 
one gonimolobe forming from the subapical cell of first-developed and second-developed 
primary gonimoblast; gonimolobes not lobed, narrow to pointed when young, becoming 
rounded when mature, carposporangia 20-35 (-40) urn in length. 

Sporangial thalli may bear tetrasporangia or bisporangia (cf. Kylin, 1907; Rosenvinge, 
1924), but usually not both, although this was reported by Harris (1959, 1962); bisporangia 
ellipsoid, 62*4 ± 5*7 x 33*4 ± 5*6 um, with transverse division; tetrasporangia sub-ellipsoid 
to sub-spheroid, 55*2 ± 4*6 x 40*9 ± 7*4 |im, with tetrahedral spore arrangement; both 
usually sessile, tetrasporangia occasionally with 1 -celled pedicel, borne adaxially on 
peripheral laterals of higher order but not prolifically on the smaller laterals of subsidiary 
axes, usually 1 (-2) per cell. 

Thalli with tetrasporangia or gametangia from March to November in the British Isles. 
The presence of tetrasporangia on gametangial plants (Feldmann-Mazoyer, 1941; Foldvik, 
1963; comments in Rueness & Rueness, 1980) or of bisporangia amongst gametangia 
(Rosenvinge, 1924) has not been detected in British field material so far. 

Ecology and distribution* 

Occurs in pools or damp sheltered areas through much of the littoral, extending to the 
sublittoral; epiphytic, commonly on Fucus, Saccorhiza, Laminaria and larger Rhodophyta. 

British Isles: Channel Islands, Cornwall, Devon, Dorset, Isle of Wight, Sussex, Kent, Norfolk, 
Yorkshire, Cheshire, Isle of Man; Caernarvonshire, Angelsey; Ayr, Bute, Argyll, Orkney, Aberdeen; 
Dublin, Cork, Galway, Mayo, Down. 

World: Eastern Atlantic from western Norway to Canaries and Morocco; western Mediterranean; 
western Atlantic from southern Massachusetts to Florida; Texas, West Indies; reported for New 
Zealand and Tasmania. 

Observations 

Callithamnion byssoides has been confused with C. corymbosum and C. roseum, but the 
species has a consistent ecological and geographical distribution. Critical cases are few in the 
British Isles, but this is not necessarily true for the western part of the north Atlantic. C. 
byssoides and C. corymbosum are occasionally confusable where the former lacks bispor- 
angia. C. byssoides usually lacks the apical pseudodichotomy at unlimited functional apices, 
the pseudodichotomies at peripheral lateral apices, and the hyaline hairs typical of C. 
corymbosum. C. corymbosum has much coarser growth and all cells are multinucleate. C. 
byssoides is finer than C. roseum, another multinucleate species. Mature cells near the 
functional apices of C. byssoides are less than 20 um maximum breadth, whereas in C. 
roseum they exceed 30 um. The often greater length of cells of C. byssoides emphasizes 
this distinction. 

The sequence of morphological phases in the life history has been established for material 
identified as Callithamnion byssoides from Port Aransas, Texas (Edwards, 1969) and from 
Wrightsville Beach, North Carolina (Kapraun, 1978). These materials may not be con- 
specific with British C. byssoides, a, point already correctly made by Rueness & Rueness 
(1980). Chromosome counts of n = 28-33 were reported in thalli from an undefined location 
in the British Isles (Harris, 1962). The recent study (Rueness & Rueness, 1980) on 
Norwegian material of C. byssoides established both a Polysiphonia-type life-history for the 

*For this and all subsequent species, distribution is stated on the basis of the old county boundaries. The 
newly-formed counties are both too large in area (particularly in Wales and Scotland) for effective discussion, and 
not yet well enough comprehended in geographical terms. 



118 P. S. DIXON & J. H. PRICE 

European taxon and an interesting mode of perennation. It is curious that although the alga 
is normally epiphytic (so reported by most authors, including this study), the perennating 
creeping filaments that subsequently gave rise to C. byssoides plants were borne on the 
ascidian Ciona intestinalis. Further field studies of this phenomenon elsewhere are 
required. 

2. Callithamnion corymbosum (Smith) Lyngbye, Tent. Hydr. Dan. : 125 (1819). 

Conferva corymbosa Smith in Smith & Sowerby, Engl. Bot.: pi. 2352 (1811). 
Ceramium versicolor C. Agardh, Syst. Alg.: 140 (1824). 

Thallus pinkish-red, or brownish to deep red in colour; to 55 mm in height; bushy, flaccid, 
usually with a single recognizable main axis; branching commonly pseudodichotomous 
throughout, but with all gradations from m©nopodial to complete pseudodichotomy; 
principal laterals of unlimited growth irregular in position and number; growth form 
irregular to more or less regular; main axes 40-270 (-300) urn in breadth; commonly 
corticate but sometimes ecorticate; occasional adventitious branching from the corticating 
filaments, but lacking formation of adventitious matrix; axial cells in straight lines, some 
curvature but no zigzags; branching often straggly and inconsistent in older parts towards 
base, younger branches show spiral pseudodichotomous arrangement, divergence angles 
usually about 30°, rarely more; peripheral laterals borne toward apices on axes of various 
orders, simple and spiral, or pseudodichotomous; apical development pseudodichotomous 
in peripheral laterals axes; apical cells of axes surpassed by peripheral laterals; apical cells of 
peripheral laterals 5-10 urn in diameter, axial apical cells often smaller; hairs apical or 
subapical, often present on all apical cells but equally often absent; cells at base of thallus 
170-250 |im in breadth, 25-35 urn in mature cells near apices; length-to-breadth ratios of 
cells 6- 1 2 to 1 ; cells multinucleate, except for a few cells near apices. 

Gametangial thalli dioecious. Spermatangiophores multi-branched, one to two cells in 
length, often several on each cell of peripheral or high order laterals; 2 to 4 spermatangia on 
ultimate cells of spermatangiophores; spermatangia 5-8 jam. 

Procarps normally formed just behind apices of principal axes and branches; usually 2 

4 

support cells, occasionally only one; carpogonial branch usually in form 4 

less often completely straight; two one-celled gonimoblasts from each central cell, 
commonly two gonimolobes on primary gonimoblast, only one on secondary gonimoblast, 
but one or more gonimolobes occasionally missing; gonimolobes rounded, usually finally 
lacking subsidiary lobing; carposporangia 25-35 urn in diameter. 

Tetrasporangia globose, 45-60 x 55-65 jim, sessile, one or more on each cell of peripheral 
laterals; often appearing to be axillary to pseudodichotomies. 

The growth form of Callithamnion corymbosum can be very variable. Thalli may possess 
either distinct axes bearing spirally arranged branches and simple peripheral laterals or with 
branches and peripheral laterals developing at same rate as principal axes, producing a 
pseudodichotomous aspect. Pseudodichotomies may occur throughout a plant, but are most 
common only in the upper parts. The number of hairs is variable; some thalli have hairs on 
almost all apical cells, others a few scattered hairs, some totally devoid of them. Harris (1959) 
noted in Galway that plants of greater shelter had a few spirally-arranged hairs in the upper 
parts while thalli of greater exposure showed pseudodichotomies and hairs on nearly all 
apical cells. This is not a difference that is consistent in such conditions (see Price, 1978). 

Thalli with gametangia reported between April and November; tetrasporangia occur 
throughout the year. 

Ecology and distribution 

Usually epiphytic, on Codium fragile, Fucus vesiculosus, or Zostera; occasionally epilithic 
in lower littoral, usually in standing water, pools and channels; known in the sublittoral to at 
least 22 m in sheltered bays. 



CALLITH AMNION IN THE BRITISH ISLES 1 1 9 

British Isles: Channel Islands, Cornwall, Devon, Dorset, Hampshire, Isle of Wight, Sussex, Somerset, 
Cheshire, Isle of Man, Suffolk, Northumberland; Pembroke, Cardigan, Merioneth, Caernarvonshire, 
Anglesey; Ayr, Bute, Argyll, Ross & Cromarty, Orkney, Aberdeen, Fife, Midlothian; Dublin, Wexford, 
Cork, Galway, Mayo, Down. 

World: Eastern Atlantic, Norway (Nordland) to Canaries; western Baltic, Mediterranean and Black 
Sea; Western Atlantic, Newfoundland to New Jersey; West Indies; Australia; Japan. 

Observations 

Callithamnion corymbosum has been confused with C byssoides and C. roseum. 
Pseudodichotomies at apices are absent in C roseum and the hairs characteristic of 
C. corymbosum are also lacking in that species. These features are also absent in C. byssoides 
but C. corymbosum has much coarser growth and multinucleate cells. 

The sequence of phases in the life history of Callithamnion corymbosum was established 
in culture for material from Helgoland (Hassinger-Huizinga, 1952). In the present study, a 
few plants have been observed with both tetrasporangia and carposporophytes and some 
with tetrasporangia and alleged bisporangia. Hassinger-Huizinga reported the occurrence in 
Helgoland material of thalli with tetrasporangia and either spermatangia or procarps in her 
cultural studies. She claimed that the expression of gametangia and tetrasporangia is 
governed by the addition or deletion of a chromosome. These data must be regarded with 
caution since neither figures nor photographs of the chromosome complements are given 
and the results necessitate absolute accuracy in counting chromosomes with a complement 
of about 60 in a nucleus 5 jim in diameter. 

In Newfoundland, Callithamnion corymbosum occurs as tetrasporangial thalli which 
appear to reproduce only by fragmentation (Whittick, 1978). 

Chromosome counts in Callithamnion corymbosum have been reported in material from 
various locations as follows: Helgoland: = 30-32; 2n = 60-64 (Hassinger-Huizinga, 1952), 
British Isles: n = 27-29 (Harris, 1959), British Isles: n = 28-33 (Harms, 1962), British Isles: 
n = c. 28 (Harris, 1966), Newfoundland: 2n = 57-65 (Whittick, 1973). 



3. Callithamnion decompositum J. Agardh, Sp. Gen. Ord. Alg. 2 (1) : 45 (185 1). 

Thallus light red in colour; somewhat rounded pyramidal in outline (see Price, 1978: 283, fig. 
15/?); rarely attaining more than 20 mm in length; main axis little corticate in British 
specimens, although dense cortication reported elsewhere; branching usually strictly 
distichous, commonly commencing at the second cell of major axes, branches restricted to 
the adaxial side of the lower two cells of primary laterals giving a slightly secund appearance 
in a few specimens; peripheral laterals attenuate and much ramified, causing flabellate 
spreading distal effect that rounds off the pyramidal outline; hyaline hair points to 60 urn in 
length reported in southern collections, but not noted in northern British material; cells of 
main axes relatively large, 130-550 \im in length, 70-200 urn in breadth in mature material; 
chloroplasts said to be polygonal-discoid when young, becoming elongate in older cells (in 
British material the variation between variable discoid and elongate does not appear to be 
correlated with age); masses of proteinaceous material present. 

Gametangial thalli unknown for the British Isles; in non-British material dioecious; 
spermatangiophores 2-4 celled, occurring on the first or second cells of laterals, each cell of 
the spermatangiophore bearing 3 or 4 spermatangia; spermatangia c. 5 x 8 urn 

Procarps formed behind apices of lateral axes, commonly with two support cells; 
carpogonial branch 'U'-shaped, trichogyne to 95um in length; gonimolobes cordate, 
41-70 x 35^47 urn; carposporangia 10x12 um in diameter. 

Tetrasporangia uncommon in British material; rounded or ovoid, to 55x65 urn in 
diameter, usually sessile, occasionally pedicellate, occurring adaxially on peripheral laterals; 
spore arrangement usually tetrahedral, occasionally irregular. 



120 P. S. DIXON & J. H. PRICE 

Parasporangia the most usual reproductive structure formed in British material, lobed or 
irregular, sometimes linear, depending on state of development, 22-56 x 1 1-22 urn, 
usually at the apices of adaxial products of peripheral lateral cells; individual paraspores 
9-18 urn. 

The form range of Callithamnion decompositum is not fully elucidated, either in the 
British Isles or elsewhere (Price, in preparation). This is partly due to uncertainties regarding 
the status of material referred by other workers to C. bipinnatum, C. tripinnatum, C. 
gallicum or C. caudatum. For a recent study of C. bipinnatum, see Rueness & Rueness 
(1980). 

Ecology and distribution 

Epiphytic on other algae, but also epizoic and epilithic; usually sublittoral, to 30 m, but also 
reported elsewhere in the lower littoral. Usually rather few thalli present. 

British Isles: Known only for Argyll (Mull), Inverness (Rhum) and Zetland, but probably occurs more 
widely. 

World: Eastern Atlantic from the Faeroes to Senegal. The species may also occur in Denmark and 
Norway, for which areas the relationships with material referred to Callithamnion bipinnatum require 
elucidation. Recent work on the latter species by Rueness & Rueness (1980) suggests that, whatever the 
application by some authors of that name, there is a good taxon concealed amongst the doubtful 
applications. 

Observations 

Callithamnion decompositum may be confused with C. hookeri in the lower littoral and 
shallow sublittoral, where both species have similar appearance and dimensions. The apical 
organization of the two species is, however, usually quite distinct with virtually absolute 
distichy in C. decompositum and some spiral arrangement of the ultimate laterals 
somewhere on the plant in C. hookeri. Below the shallow sublittoral, C. hookeri becomes 
more lax and less branched, whereas C. decompositum retains its characteristic growth-form. 

Although parasporangia are the most reproductive structures found in British material, 
there is insufficient information to comment on their phenology. 

Nothing is known of the life history of Callithamnion decompositum. These northern 
populations are parasporangial, although gametangia and/or tetrasporangia are produced by 
material from areas south of the British Isles. In Finistere, parasporangia have been reported 
mixed with tetrasporangia. Data on the life history in northern material will, it is hoped, be 
more fully available when the Faeroe Islands specimens at present in culture at the 
University of Oslo (J. and M. Rueness) have developed beyond vegetative growth from the 
originally-released tetrasporangia. 

4. Callithamnion granulatum (Ducluzeau) C. Agardh, Sp. Alg. 2(1): 177 (1828). 

Ceramium granulatum Ducluzeau, Essai Hist. Nat. Conf. Montpellier: 72 (1806). 
Callithamnion harveyanum J. Agardh in Linnaea 15 : 45 (1841). 
Callithamnion grande J. Agardh, Alg. Mar. Med. Adr:. 73 (1 842). 

Thallus pale straw, brownish, rust, to purple-red or dark red in colour; to 1 50 mm in height; 
robust but highly flexible, flaccid in water; spongy in texture and appearance, but less so than 
in C. sepositum; appearing as regular or irregular, simple or compound pine-tree like 
pyramids; main axes rarely bifurcate near base, to 3 mm or more in breadth, but probably 
greater in larger specimens; strongly corticate throughout; cortication develops dense matrix 
of short tightly-woven adventitious laterals, giving rise occasionally to peripheral laterals of 
normal form; main axial cells almost invisible without dissection; axial cells in straight lines, 
length-to-breadth ratios of any visible upper axial cells usually 1*5 to 3:1; branching 
monopodial, associated with pseudo-dichotomy at all levels and especially obvious at 



CALLITHAMNION IN THE BRITISH ISLES 121 

peripheries of thallus; divergence angles greater than 45°, sometimes 90°, unless strongly 
appressed in corymbose clusters; apical cells of indeterminate laterals buried in the 
corymbose clusters of peripheral laterals, normally needing dissection for examination; 
peripheral laterals often spreading, but also frequently grouped in corymbose clusters, that 
represent individual lateral systems, to 700 urn in breadth (Price, 1978 : Fig. 11A-C); 
individual peripheral laterals often straight although occasionally curved in the corymbose 
clusters; greatest width of peripheral lateral at or near base; cells (1-) 2-4 (-6) per peripheral 
lateral, usually more or less rectangular, with some apical narrowing; peripheral lateral 
apical cell a short cylinder to roundly-truncated cone, 1 7*6 ± 6*6 um in length; with 
occasional terminal or subterminal hyaline hairs. 

Gametangial thalli dioecious or (less often) monoecious. Spermatangial groups hemi- 
spherical, rarely coalescent; spermatangiophores multibranched, branches 3-5 cells long; 
two to four spermatangia on peripheral cell of each branch; spermatangia ovate to ellipsoid, 
6x8 um. 

Procarps formed just behind functional apices of penultimate and lower order laterals; 
usually two support cells, but occasionally one; carpogonial branch form difficult to 
interpret, but straight, with minor variations, and zigzag (the more usual) noted; carpo- 
gonial maximum breadth 18 urn; trichogyne to 70-240 jam in length; gonimoblasts single- 
celled, two from each central cell; usually one gonimoblast giving rise to one gonimolobe, 
occasionally gonimolobes missing; gonimolobes rounded without subsidiary lobing: carpo- 
sporangia 25-35 |im in diameter. 

Tetrasporangia initially apical on cell below peripheral dichotomies, spreading to cells of 
peripheral laterals, large, globose to ellipsoid, 45-60 x 50-60 um, never more than one per 
bearing cell; polysporangia reported but never observed in the present study. 

Detailed consideration of some aspects of growth form variation in Callithamnion 
granulatum is given in Price (1978). 

Reproductive phenology in Callithamnion granulatum is variable with location, year and 
individual thalli, although patterns can be perceived. Detailed information is being 
published elsewhere. 

Ecology and distribution 

Occurs in rock pools and on emersed damp surfaces in the lower littoral of wave-exposed 
shores; commonly epiphytic, very rarely epilithic, frequently epizoic on Mytilus, Patella and 
Balanus, especially in blanket populations; reported from the sublittoral, but all such reports 
erroneous except in very shallow conditions (to 0*5 m). 

British Isles: Channel Islands, Cornwall, Devon, Dorset, Isle of Wight, Yorkshire, Isle of Man; 
Pembroke, Caernarvonshire, Anglesey; Ayr, Bute, Argyll, Inverness, Ross & Cromarty, Orkney, 
Zetland, Angus, Fife, Midlothian, East Lothian; Antrim, Mayo, Galway, Clare, Cork, Wexford, 
Wicklow, Dublin, Down. 

World: North-eastern Atlantic from Shetlands to Morocco, perhaps to the Gulf of Guinea, locally 
(Liberia, Cote d'lvoire, Ghana); Mediterranean. Material reported from the Faeroes all appears 
referable to the exposed-water form o{ Callithamnion hookeri, growing epiphytically on C sepositum. 

Observations 

The only possible confusions are with Callithamnion sepositum or C spongiosum. The 
principal criterion for discrimination is the most usual number of cells in the highest order 
peripheral laterals, with 1 or 2 cells in C spongiosum, 3 to 4 in C. granulatum and 7 to 9 in 
C sepositum. C granulatum may also be distinguished from C sepositum in its consistent 
pseudodichotomy and the basal, or near basal, maximum width of the peripheral laterals. 
The greatest difficulty is in discriminating from C spongiosum, with which it tends to occur 
in mixed populations although intermediate forms are lacking. Cotton (1912) distinguished 



122 P. S. DIXON & J. H. PRICE 

between the two species, although the ecological differences which he noted do not hold 
consistently. 

Little published information is available on the life history of Callithamnion granulatum; 
all exposed shore forms are difficult to culture. Harris (1962) reported a chromosome 
number of n = 28-33 in material identified by him as C. purpurascens and possibly referable 
to this entity. 



5. Callithamnion hookeri (Dillwyn) S. F. Gray, Nat. An. Br. PL 1 : 324 (1821). 

Conferva hookeri Dillwyn, Br. Conf.: pi. 106 (1809). 
Callithamnion roseum /? tenue Lyngbye, Tent. Hydr. Dan.: 126 (1819). 
Ceramium scopulorum C. Agardh, Syst. Alg.\ 132 (1824). 
Callithamnion scopulorum C. Agardh, Sp. Alg. 2(1): 166 (1828). 
Callithamnion poly spermum C. Agardh, Sp. Alg. 2(1): 169 (1828). 
Callithamnion brodiaei Harvey in Hooker, Eng. Fl. Sir J. E. Smith 5 : 340 (1 833). 
Callithamnion grevillei Harvey in Hooker, Eng. Fl. Sir J. E. Smith 5 : 345 (1833). 
Callithamnion spinosum Harvey in Hooker, Eng. Fl. Sir J. E. Smith 5 : 345 (1833). 
Callithamnion acrospermum J. Agardh, Sp. Gen. Ord. Alg. 2(1): 52 (185 1). 

Thallus of widely different colours but usually dark to brownish red; to 70 mm or more in 
height; bushy, straggly, or sparsely branched, sometimes even reduced to a single main axis; 
main axis or axes 300-500 jim in breadth, commonly corticate, often with reduced 
cortication, or even none in younger material; cortication often producing adventitious 
laterals, when extreme forming a matrix that may be over 2 mm overall breadth; branching 
very variable, from regular alternate-spiral to highly irregular; peripheral laterals distichous 
or spiral, but not usually overtopping the apex of main axis, borne on penultimate laterals 
with strong axial zigzag; apical cells rounded, basal breadth 5-10 urn; cell numbers and 
dimensions in peripheral laterals highly variable; cells of peripheral laterals cylindrical or 
slightly tapered; mature cells near base of thallus 50 to 100 urn maximum diameter, reducing 
to 40 to 80 urn near apex, length-to-breadth ratios varying between 0*5 to 1 and 5 to 1 . 

Gametangial thalli dioecious. Spermatangiophores one, rarely more, per cell of peripheral 
laterals, variable in form even in same thallus; most spermatangiophores consisting of a 
single cell with a single secund branchlet, 3 to 4 cells in length, others with a series of secund 
branchlets; ultimate cells of branchlets bearing 2 to 4 spermatangia, usually secund but 
sometimes distichous; spermatangia 4x8 um. 

Procarps formed just behind apices of axes and indeterminate laterals; carpogonial branch 
of variable form, from straight to zigzag; usually developing two auxiliary cells after 
fertilization, but occasionally only one; each auxiliary cell producing two single-celled 
gonimoblasts, with one or two gonimolobes, although the second-developed filament may 
produce only one or even none; gonimolobes lobed or simple, usually rounded, but 
sometimes pointed or even conical; carposporangia at maturity 35-50 um in diameter. 

Tetrasporangia globose or nearly so, sessile, 40-60 x 45-60 urn, borne adaxially on 
peripheral laterals, usually one per cell, but occasionally with a second tetrasporangium 
borne abaxially. Bisporangia occur rarely, mixed with tetrasporangia, and may represent 
only an aberrant or developmental stage of the latter. Parasporangia replacing vegetative 
lateral filaments reported widely, but not authenticated in the British Isles. Monosporangia 
reported, but probably nothing more than tetrasporangial initials. 

The overall thallus form is highly variable under different environmental conditions and 
in different locations. So far as can be determined, there are no consistent morphological 
distinctions between Callithamnion hookeri and the entities usually treated under the names 
C. scopulorum, C. brodiaei and C. polyspermum. The thalli which have been referred to C. 
scopulorum occur in the upper littoral in damp locations as clumps or local turfs. The 
customary epiphytic littoral populations tend to be lax whereas in situations exposed to 
wave-action the thalli are robust, short, stubby and much branched. The sublittoral 



CALLITH AMNION IN THE BRITISH ISLES 1 23 

populations tend to be extremely lax. Analysis of this variation in respect to previous 
nomenclature and taxonomy is given in Price (1978). 

Reproductive phenology in Callithamnion hookeri is variable with location, year and 
individual thalli, although patterns can be perceived. Detailed information is being 
published elsewhere. 

Ecology and distribution 

Occurs in rock pools and on emersed damp surfaces in the lower littoral of wave-exposed 
shores; commonly epiphytic, rather rarely epilithic, frequently epizoic on Mytilus, Patella 
and Balanus. 

British Isles: Callithamnion hookeri is probably present throughout the British Isles, occurring under 
appropriate circumstances. Positive identifications are, however, available only for the following 
counties; Channel Islands, Cornwall, Devon, Dorset, Hampshire, Sussex, Kent, Essex, Suffolk, 
Norfolk, Lincolnshire, Yorkshire, Durham, Northumberland, Somerset, Cheshire, Cumberland, Isle of 
Man; Glamorgan, Pembroke, Cardigan, Caernarvonshire, Anglesey; Dumbarton, Wigtown, Ayr, 
Bute, Renfrew, Argyll, Inverness, Ross & Cromarty, Orkney, Zetland, Sutherland, Moray, Aberdeen, 
Kincardine, Angus, Fife, Midlothian, East Lothian, Berwick; Antrim, Londonderry, Donegal, Leitrim, 
Mayo, Galway, Clare, Limerick, Kerry, Cork, Waterford, Wexford, Wicklow, Dublin, Down. 

World: Callithamnion hookeri is extremely widespread but a precise statement of its total distribution 
is difficult in view of its great variability and the need to clarify the status of material referred to such 
entities as C. bipinnatum, C tripinnatum, C. pusillum, C. hirtellum, C truncatum, etc. The species 
occurs in the north Atlantic in Iceland and between northern Norway and the Canary Islands and 
Morocco, occurring also (Price, in press) in areas of cold upwelling in the Gulf of Guinea 
(and perhaps sparsely in Angola), on the Baltic Sea coast of Germany, and from Newfoundland (South 
& Hooper, 1980) southward. The reported occurrence of various entities possibly referable to this 
species in the Mediterranean must await taxonomic clarification; the same comment applies to reports 
from the Pacific Ocean. 

Observations 

The life history of Callithamnion hookeri has been shown by culture studies of British 
material to consist of a sequence of gametangial, carposporangial and tetrasporangial 
phases (Edwards, 1979), although nothing is known of the chromosome counts. In 
populations from the west coast of Sweden, it has been shown that paraspores develop into 
parasporangium-bearing thalli for several successive generations (Rueness & Rueness, 1978). 
The role of the alleged monosporangia has not been established. On the relations between 
life-history, tolerance levels, and distribution pattern, see van den Hoek (in press). 

6. Callithamnion roseum (Roth) Lyngbye, Tent. Hydr. Dan.: 126, tab. 39 (1819). 

Ceramium roseum Roth, Archiv Bot. (Romer) 1 (3): 47 (1798). 

Conferva rosea (Roth) Smith in Smith & Sowerby, Engl. Bot.: pi. 966 (1802). 

Callithamnion octosporum C. Agardh, Sp. Alg. 2(1): 177 (1828). 

Thallus pinkish to purple-red; to 80 or more mm in height; bushy and fairly robust, less 
delicate or flouncy than Callithamnion byssoides or C. corymbosum, but still flaccid; usually 
with one main axis; main axis to 1-0 mm in breadth, sometimes ecorticate but commonly 
with at least a few cortical filaments from lower branches, occasionally moderately corticate 
but no development of adventitious matrix from cortication; branching monopodial but 
sometimes with pseudodichotomous aspect below carposporophyte; axial (indeterminate) 
branches irregular in position and number, variable in length, especially at base of plant, 
spirally arranged in upper parts; peripheral laterals usually simple, upsweeping and 
incurving, spiral; apical cells of peripheral laterals 5-10 urn in breadth; apical cells of main 
axis and main branches surpassed by subordinate peripheral laterals; cells at bases of plants 
130-225 jim in breadth; tapering; mature cells, near apices, 30-35 urn in breadth; length-to- 
breadth ratio 5-12 to 1 but cells usually shorter, more robust, than C. byssoides; cells usually 



124 P. S. DIXON & J. H. PRICE 

multinucleate except immediately behind apices; chloroplasts irregularly discoid in young 
cells, forming continuous ribbons with age. 

Gametangial thalli dioecious. Spermatangiophores two cells in length, multibranched, 
often several on each cell of peripheral laterals; two to four spermatangia on ultimate cells of 
spermatangiophore cluster; spermatangia 5x8 um. 

Procarps formed behind apices of main axes and branches, commonly with two support 
cells, rarely only one; carpogonial branch straight, occasionally zigzag; two gonimolobes 
from each central cell, each one-celled; commonly two gonimolobes on first gonimoblast, 
but only one on secondary gonimoblast; one or more gonimolobes occasionally missing; 
gonimolobes usually lacking subsidiary lobing, rounded; carposporangia 20-35 urn in 
diameter. 

Tetrasporangia globose, 40-50 x 45-55 um, sessile, often abundant, one or more, usually 
adaxial, on each mature cell of peripheral laterals. 

Spermatangial thalli of Callithamnion roseum tend to be more delicate than tetra- 
sporangial or sterile specimens, but the degree of robustness varies enormously. 

Fertile thalli appear to occur at all times of the year, but there is considerable variation in 
the reproductive organs present according to individual thallus, year, time and location. 

Ecology and distribution 

Epiphytic on other algae in lower littoral in sheltered conditions or sheltered niches on more 
wave-exposed shores; can extend into the sublittoral to at least 20 m. A characteristic habitat 
is just below water level on floating structures in areas of somewhat fluctuating salinity. 

British Isles: Channel Islands, Cornwall, Devon, Hampshire, Sussex, Kent, Suffolk, Norfolk, 
Lincolnshire, Isle of Man; Caernarvonshire, Anglesey; Cork, Galway, Wexford, Dublin. Probably 
widespread. 

World: Eastern Atlantic from Norway to Portugal; Baltic; eastern coast of the United States. 

Observations 

Callithamnion roseum may be confused with C corymbosum and C byssoides, but lacks 
pseudodichotomous apical development and the hairs which occur quite commonly in 
British C corymbosum. C roseum has coarser growth than C byssoides, with multinucleate 
cells. 

Tetrasporangia occur in all months of year in most areas of the British Isles. Gametangial 
plants and carposporophytes detected April-November. Little published information exists 
on the life history of Callithamnion roseum. Bisporangia or carposporophytes have been 
reported on a few tetrasporangial plants. Chromosome number reported, doubtfully, as n = c. 
39 (Harris, 1962, 1966), but no other data available. 



7. Callithamnion sepositum (Gunnerus) P. Dixon & J. Price, comb. nov. 

Conferva seposita Gunnerus, Fl. Norv.: 1 1 6 (1 772). 
Conferva purpurascens Hudson, Fl. Angl., 2nd ed. 2 : 600 (1778). 
Conferva arbuscula Dillwyn, Br. Conf.: pi. 85 (1807). 
Ellisius glaber S. F. Gray, Nat. Arr. Br. PL 1 : 333 (182 1). 
Phlebothamnion faroense Kiitzing, Tab. Phyc. 14 : 30 (1864). 
Phlebothamnion scoticum Kiitzing, Tab. Phyc. 14 : 30 (1864). 

Thallus dark red to dark purple in colour, distally bleaching to straw-coloured; robust and 
spongy but highly flexible, the texture resembling wet wool; main axes and laterals strongly 
corticate, covered throughout by a matrix of short, adventitious laterals obscuring the axes; 
axes from 310 urn to more than 5 mm in breadth; branching alternate-pinnate throughout, 
with occasional apparent aberrant pseudodichotomies; intermediate laterals arising in all 
directions; peripheral laterals grouped into tightly-packed, incurved, corymbose clusters, 



CALLITH AMNION IN THE BRITISH ISLES 1 25 

700 to 1400 jim in breadth (Price, 1978 : Fig. 10A-C); apical cells truncated cones, length 
frequently greater than breadth, length 20*5 ± 5*0 jxm, basal breadth 1660 ± 4*2 urn, 
apical breadth 5'50 ± 2*5 jam; cell numbers in peripheral laterals (2-) 7-9 (-16); peripheral 
laterals 125-539 jim in length x 24-60 |um in breadth, position of the maximum breadth 
usually well above the base or lower half parallel-sided; cells of peripheral laterals 
cylindrical, the maximum breadth equal to the length. 

Gametangial thalli monecious or dioecious, predominance apparently varying with 
region. Spermatangial thalli with dense spermatangia within the apical corymbose clusters, 
sometimes in localized areas, sometimes widespread throughout; initial stages adaxial along 
incurved peripheral laterals, but subsequently spreading to form coalescent cushions along 
the lengths of whole laterals. 

Carpogonial thalli usually bearing abundant procarps; position of procarps variable, 
above or below outer forkings on peripheral laterals in heavy-cropped specimens; two 
support cells developed, frequently only one visible; carpogonial branch apparently 
U-shaped but sometimes with possible zigzag; carpogonia 15-26 jim in breadth, trichogyne 
to 240 urn in length, always twisted, occasionally with swellings or bifurcations; carpo- 
sporophytes large and conspicuous when fully grown, normally paired on opposing support 
cells, occasionally unilateral; in luxuriant growth, carposporangia becoming angular through 
lateral pressure; rounded-off carposporangia 70 x 70 jim. 

Tetrasporangia sub-spherical to spherical, sessile; (42-) 59-77 (-110)x(37-) 55-70 
(-100) jim; adaxial, occasionally abaxial on cells of outer peripheral laterals. 

There are correlations between the form of mature plants and reproductive state. 
Tetrasporangial specimens show luxuriant clumped and bushy branching, while carpo- 
sporangial thalli show sparser, more open, branching with longer laterals, which however do 
not contain more cells. Spermatangial specimens are intermediate between these two. 
Hyaline hair points can occur on any material but particularly on carpogonial plants. 
During winter and early spring, thalli of Callithamnion sepositum are very straggly and tend 
to lack the peripheral corymbose clusters of laterals. 

Reproductive phenology in Callithamnion sepositum is variable with location, year and 
individual thalli, but patterns can be perceived. Detailed information is being published 
elsewhere. 



Ecology and distribution 

Commonly epilithic on steep slopes or epizoic on shells of Patella, Balanus and Mytilus; in 
blanket populations, occasionally epiphytic on Corallina or on the bases of exposed Fucus 
vesiculosus; very rarely in pools except in blanket populations. 

British Isles: Cornwall, Devon, Yorkshire, Durham, Northumberland, Isle of Man; Pembroke, 
Caernarvonshire, Anglesey; Wigton, Ayr, Bute, Argyll, Inverness, Ross & Cromarty, Sutherland, 
Caithness, Orkney, Zetland, Banff, Aberdeen, Angus, Fife, Midlothian, East Lothian, Berwick; 
Waterford, Cork, Kerry, Clare, Galway, Mayo, Donegal. 

World: Norway, Faeroes, and Iceland. 



Observations 

Callithamnion sepositum is similar in size, appearance and growth in blanket populations 
on the shore to C granulatum, but mature thalli of C. sepositum lack the symmetrical 
appearance of those of C granulatum. See pp. 121-2; 126-7 where the differences are 
discussed in detail. 

Little published information is available on the life history of Callithamnion sepositum; 
all exposed shore forms are difficult to culture. Harris (1962) reported a chromosome 
number of n = 28-33 in C. purpurascens, possibly referable to this entity. 



126 P. S. DIXON & J. H. PRICE 

8. Callithamnion spongiosum Harvey in Hooker, Engl Fl. Sir J. E. Smith 5 : 346 (1833). 

Thallus through pale straw or brownish or rust, to purple-red or dark red in colour according 
to season and environment; to 50 mm in height; robust but highly flexible, flaccid in water; 
spongy in texture and appearance, but (like Callithamnion granulatum) less so than in C. 
sepositum; appearing as regular or irregular, simple or compound pine-tree like pyramids; 
main axes rarely bifurcate near base; to 3*0 mm in breadth, but probably greater in larger 
specimens; strongly corticate throughout; cortication developing dense matrix of short 
tightly woven adventitious laterals, giving rise occasionally to peripheral laterals of normal 
form; main axial cells almost invisible without dissection; axial cells in straight lines, length- 
to-breadth ratios of any visible upper axial cells usually 1*5-3 to 1; branching monopodial, 
associated with pseudodichotomy at all levels but especially obvious at periphery of thallus; 
divergence angles greater than 45°, sometimes 90°; indeterminate apical cells buried in 
corymbose clusters (representing individual lateral systems) of peripheral laterals, normally 
needing dissection for examination; peripheral laterals often spreading, frequently grouped 
in corymbose clusters to 700 urn in breadth; individual peripheral laterals usually straight 
(although occasionally curved in corymbose clusters) consisting of one, less often two, cells, 
greatest width at or near base; cells usually more or less cylindrical, with some apical 
narrowing; apical cell short cylinder or rounded, 1 76 ± 6*6 jim in length; occasional 
terminal or subterminal hyaline hairs. 

Gametangial thalli dioecious or monoecious. Spermatangial cluster hemispherical, 
rarely coalescent; spermatangiophores multibranched, branches 3-5 cells long; two to four 
spermatangia on peripheral cell of each branch; spermatangia ovate to ellipsoid, 6x8 urn 

Procarps formed just behind functional apices of penultimate and lower order laterals; 
usually two support cells, but occasionally one; carpogonial branch form difficult to 
interpret, but straight (with minor variations) and zigzag noted; carpogonial maximum 
breadth 18 urn, trichogyne to 70-240 jxm in length; gonimoblasts single-celled, two from 
each central cell; usually one gonimoblast giving rise to one gonimolobe, but occasionally 
gonimolobes missing, mature gonimolobes rounded without subsidiary lobing; carpospor- 
angia 25-35 urn in diameter. 

Tetrasporangia initially apical on cell below peripheral dichotomies, spreading to cells of 
peripheral laterals, large globose to ellipsoid, 45-60 x 50-60 jum, never more than one per 
bearing cell; polysporangia reported but never observed in the present study. 

So far as can be determined from the sparse material available, the growth form variation 
in Callithamnion spongiosum is similar to that of C granulatum, discussed on p. 120 et seq. 

Callithamnion spongiosum occurs too sparsely for any meaningful comments to be made 
on the reproductive phenology of this species. 

Ecology and distribution 

Occurs in rock pools and on emersed damp surfaces in the lower littoral, commonly 
epiphytic, less frequently epizoic on Mytilus, Patella and Balanus. Very rarely in large 
populations; usually mixed as sporadic thalli with Callithamnion granulatum. 

British Isles: Cornwall, Devon, Isle of Man; Pembroke, Caernarvonshire, Anglesey; Ayr, Bute; 
Antrim, Dublin. 

Other records (Sussex, Yorkshire, Fife, Zetland, Angus, Channel Islands) have proved on 
re-examination to be erroneous, although there is no reason to doubt the sporadic occurrence of the 
entity in western areas. 

World: No verifiable records are known, but on the basis of present knowledge the presence of this 
species in the least north-west France is probable. 

Observations 

The only possible confusions are with Callithamnion sepositum or C granulatum. The most 
usual number of cells in the higher order peripheral laterals is the prime characteristic for 



CALLITH AMNION IN THE BRITISH ISLES 127 

discrimination, with 1 or 2 cells in C. spongiosum, 3 to 4 in C. granulatum and 7 to 9 in C. 
sepositum. C. spongiosum may also be distinguished from C. sepositum in its consistent 
pseudodichotomy and the basal maximum width of the peripheral laterals. The greatest 
difficulty is in discriminating from C. granulatum, with which it tends to occur in mixed 
populations, although intermediate forms are lacking. Cotton (1912) distinguished between 
the two species, although the ecological differences which he noted do not consistently hold. 

It may be that the rarity of occurrence of specimens amongst Callithamnion granulatum 
populations represents the detected manifestation of tendencies towards speciation in a 
genome currently still interfertile with the C. granulatum from which it arises. If this is so, 
the expected intermediates must be being dismissed as plants of C. granulatum. The 
inconsistency and sparseness of appearance in particular C. granulatum populations may 
indicate a present lack of success in establishment as a separate and defined breeding 
population (eventually, species). It will be interesting to see if specimens of C. spongiosum 
become more frequent in the field in the (imponderable) future, with some gradual emphasis 
of ecological tolerance differences, at present not recognizable to us. 

No published information is available on the life history o$ Callithamnion spongiosum', all 
exposed shore forms are difficult to culture. Harris (1962) reported a chromosome number of 
n = 28-33 in material referred by him to C. purpurascens, possibly referable to this entity. 

9. Callithamnion tetragonum (Withering) S. F. Gray, Nat. An. Br. PL 1 : 329 (1821). 

Conferva tetragona Withering, An. Br. PL, 3rd ed. 4 : 405 (1 796). 

Ceramiumfruticulosum Roth, Cat. Bot. 2 : 183 (1800). 

Conferva fruticulosa Wulfen, Crypt. Aquat. : 26 (1803). 

Ceramium brachiatum Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 136 (1828). 

Ceramium dudresnayi Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 125 (1828). 

Ceramium guttatum Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 127 (1828). 

Callithamnion fruticulosum J. Agardh in Linnaea 15 : 46 (1841). 

Thallus red, brownish-red to rust in colour; robust, coarser than all other species except 
Callithamnion tetricum, irregularly pyramidal in outline, to 110 mm in height; usually a 
single main axis in a thallus, apparent basal division into several principal axes often 
resulting from separate development of adjacent spores; branching regularly spiral, 
occasionally irregular towards base of main axis, larger laterals similar to main axis; main 
axis strongly corticate, 233 to 550 |im (to 1 mm) in breadth; adventitious branching from 
the cortication absent or sparse; axial cells in linear rows, rarely even distal axial zigzag; 
length-to-breadth ratios of axial cells easily seen through cortication, usually less than 3 to 1 ; 
cell breadth tapering from axial base to apex, from 200-300 to 90-1 10 urn; apices surpassed 
by clustered peripheral laterals, latter without the corymbose grouping that occurs in many 
other exposed shore species; older lower order laterals easily confused with main axes when 
detached; peripheral laterals incurved, often strongly so giving 4 awl'-shape, occasionally 
straighter, longer and narrower, appearing almost parallel-sided ('brachiatum'); cells of 
peripheral laterals barrel-shaped ('tetragonum') or cylindrical (' brachiatum'); apical cells of 
peripheral laterals small in relation to products, more or less conical, with pointed apices, 
12 ± 3 urn, in length, 10 ± 4 urn in breadth at base; cells below determinate lateral apices 
truncated cones; breadth of immediate derivatives equal to or greater than length; cells 
usually two or more times as long as apical cell; cells in peripheral laterals 1 50 to 700 urn in 
length, maximum breadth, basal to median in position, (11-) 40-165 urn; hairs absent from 
British material; cells multinucleate, except near apices; chloroplasts irregularly discoid, 
occasionally becoming long thin ribbons or irregular masses. 

Gametangial thalli usually monoecious, habitually so in some populations. Spermatangio- 
phores many-branched, branches 3 to 5 cells long, forming unilateral adaxial tufts, 
60 ± 6|im in length, 30 ± 7 jim in height, one per cell of peripheral laterals, borne near 
apical septum, rarely coalescent; ultimate cells bearing two to four spermatangia per cell; 
spermatangia 5x8 urn 



128 P. S. DIXON & J. H. PRICE 

Procarps formed behind indeterminate apices of peripheral branch systems, often basal to 
clusters of peripheral laterals; normally two support cells, occasionally only one; 
carpogonial branch of variable form, but usually straight; trichogyne shape and size variable, 
with various swellings and usually divided distally, 50 to 200 um in length; each auxiliary 
cell producing two single-celled gonimoblasts and a single gonimolobe from each gonimo- 
blast cell; mature gonimolobes rounded, not lobed; carposporangia variable in shape, due to 
appression, 25-45 um in diameter. 

Tetrasporangia spherical to ellipsoid, sessile, 60 ± 8 x 52 ± 7 um, borne adaxially on 
peripheral laterals, rarely more than one per cell. 

The overall thallus form of Callithamnion tetragonum is usually pyramidal and quite 
characteristic, only rarely appearing straggly and irregular. The peripheral laterals exhibit 
varying degrees of curvature and cell form, the extremes having been sufficiently distinct for 
two species to have been recognized formerly, although now treated merely as forms. In 
some thalli, the laterals are strongly incurved {tetragonum form; Price, 1978 : Fig. 13), whilst 
in others they are rather straighter and much narrower (brachiatum form; Price, 1978 : Fig. 
14A): mixtures of the two types are also known. 

Reproductive phenology in Callithamnion tetragonum is variable with location, year and 
individual thalli, but patterns can be perceived. Detailed information is being published 
elsewhere. 

Ecology and distribution 

Occurs in rock pools or open damp conditions at all levels, except high littoral and supra- 
littoral, but best developed in the lower littoral under conditions of considerable, but not 
extreme, wave action, and in the sublittoral to 24 m; predominantly epiphytic, more rarely 
epilithic. Usually more abundant and luxuriant from the shallow sublittoral, especially on 
the laminae of Laminaria spp. 

British Isles: Channel Islands, Cornwall, Devon, Somerset, Dorset, Hampshire, Isle of Wight, Sussex, 
Kent, Yorkshire, Durham, Northumberland, Isle of Man; Glamorgan, Pembroke, Cardigan, 
Caernarvonshire, Anglesey, Denbigh; Bute, Argyll, Ross & Cromarty, Orkney, Aberdeen, Angus, Fife, 
East Lothian, Berwick; Derry, Down, Antrim, Dublin, Wexford, Waterford, Cork, Kerry, Clare, 
Galway, Mayo. 

World: North-eastern Atlantic from Iceland and Norway to the Algarve; Mediterranean; eastern coast 
of U.S.A. Recorded, possibly in error, from the Canaries, Cape Verde Islands, and the Cape of Good 
Hope. 

Observations 

Callithamnion tetragonum is not easy to mistake; even so, there have been numerous 
misidentifications. 

Little information is available on the life history of Callithamnion tetragonum. Life 
history of the American C baileyi, considered by some to be conspecific with C tetragonum, 
has recently been elucidated by Whittick & West (1979). Chromosome counts n = 9-10, 
2n= 18-20 (Mathias 1927, 1928, 1932, 1935) were shown to be erroneous by Westbrook 
(1930a, 1935); n = 28-33 (Harris, 1962). 



10. Callithamnion tetricum (Dillwyn) S. F.Gray, Nat. Arr. Br. PI. 1 : 324(1821). 

Conferva tetrica Dillwyn, Br. Conf. : pi. 81 (1806). 

Ceramium congestum Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 125 (1828). 

Thallus red to brown in colour, rarely purple; rope-like, coarse and straggly, to 250 mm in 
height; main axes and major laterals densely corticate, cortication giving rise to very thick 
matrix of adventitious laterals; main axes 2-4 mm in breadth; branching strictly mono- 
podia!, indeterminate branches irregular in position and development, resembling main 



CALLITHAMNION IN THE BRITISH ISLES 129 

axes; distal middle order laterals bearing stiff, shorter but progressively lengthening 
peripheral laterals giving a 'conical' (narrow pyramidal) outline to the individual branching 
system, the lower few cells of these distal and larger peripheral laterals occasionally bare; 
higher (later) order laterals distichous, angle of insertion acute, 50-60°; apical cells of bearing 
indeterminate laterals never over-topped by peripheral laterals (see Price, 1978 : Fig. 12); 
apical cells 125-175 fim near base of thallus, 75-100 jim behind distal apices of thallus; cell 
length-to-breadth ratios 1-3 (-5) to 1 in apical parts, (1-) 3-5 to 1 in basal parts of thallus; 
cells lengths and breadths increasing rapidly for about 4 to 6 cells behind the apical cell and 
immediate derivatives; chloroplasts irregular, discoid, or ribbon-shaped. 

Gametangial phase dioecious. Spermatangiophores branched, branches 3-5 celled, borne 
on every cell of peripheral lateral, in luxuriant development forming large coalescent masses; 
2-4 spermatangia from ultimate cells of spermatangiophores; spermatangia 4-8 p.m. 

Procarps formed behind apices, especially of peripheral laterals; usually two bilateral 
support cells, occasionally only one; carpogonial branch straight, less often zigzag; each 
central cell developing two 1 -celled gonimoblast filaments which bear 1 gonimolobe each; 
mature gonimolobes with no subsidiary lobing; carposporangia 8-12 jim in diameter. 

Tetrasporangia globose, 65-90 x 75-90 |im, sessile, adaxial on peripheral laterals, one per 
cell; bi- and tri-sporangia reported (Westbrook, 1930). 

Habit varies little. Specimens more straggly over the winter and branching sparser and 
more open in sublittoral material. Perennial at least at times in Britain and probably 
throughout range. Branching pattern correlates with reproductive state, but seasonal effects 
probably equally as great. 

Reproductive phenology in Callithamnion tetricum is variable with location, year and 
individual thalli, but patterns can be perceived. Detailed information is being published 
elsewhere. 

Ecology and distribution 

Usually epilithic or less commonly epiphytic on shores exposed to strong wave-beat, but 
never on the fully exposed positions that encourage especially Callithamnion sepositum and 
to a lesser extent C granulatum on very exposed shores; C. tetricum occurs only in shore 
conditions offering some protection from direct wave impact (see Price, 1978). 

British Isles: Channel Islands, Cornwall, Devon, Dorset, Hampshire (Isle of Wight), Kent, Norfolk 
(doubtful), Isle of Man; Glamorgan, Pembroke, Anglesey; Down, Dublin, Wexford, Waterford, Cork, 
Clare, Galway. 

World: Known from north Ireland to Morocco. 

Observations 

Callithamnion tetricum, wherever it occurs, is one of the most distinctive species of the genus 
and has been involved in few confusions. 

Little is known of the life history of Callithamnion tetricum. Reported chromosome 
numbers show curious disagreements. Westbrook (1930/?) reported ?n = c.25 and ?2n = c.50, 
while Harris (1959) reported counts of 92, 93, 94 and 98 in sporangia, later cited (Harris, 
1962) as '7 90-100'. The latter counts are three times greater than those obtained in any 
other species of Callithamnion and approximately twice the number reported by Westbrook. 



Rejected species 
1. Rejections from the British flora 

Two species of Callithamnion accepted in the British flora for many years must be rejected, 
either on the grounds oftaxonomic uncertainty or because of misidentifications. These are: 



1 30 P. S. DIXON & J. H. PRICE 

Callithamnion rabenhorstii (Kiitzing) P. Crouan & H. Crouan. 

Described originally from the eastern Mediterranean (Ionian Sea) by Kiitzing (1849 : 896), 
the species was added to the British flora on the basis of material collected by E. M. Holmes 
at Studland (Dorset). The status of the entity is questionable, even at its original location, and 
there is no valid reason to accept the species in the British flora. Harris (1966) has suggested 
that the Holmes material resembles C. byssoides, and this attribution appears to be correct. 

Callithamnion tripinnatum C. Agardh 

This entity has been accepted in the British flora for many years, although it has been the 
subject of much confusion. The initial record for the British Isles, bv Harvey (in Hooker, 
1833 : 346), is based on material of Compsothamnion sp. Subsequently, Harvey (1847, pi. 
LXXVII) corrected this initial error, although the plants to which the name was then applied 
are referable to Callithamnion hookeri. Whatever the status of the original material of C. 
tripinnatum might be, as stated on p. 123 it would seem to form part of the C hookeri 
assemblage; there is no valid reason for accepting C. tripinnatum in the British flora. 

2. Entities of uncertain attribution 

Several species, attributable to the genus Callithamnion and described from the shores of the 
British Isles and adjacent regions, cannot be referred with certainty to any of the previously 
mentioned species. There are two principal reasons for this uncertainty: 

1 . The type material cannot now be located, if it ever was preserved. 

2. The type material has been located, but is inadequate in quality or quantity for 
determination. 

The entities of uncertain attribution are: 
Callithamnion arachnoideum Griffiths ex Moore in Larcom, Ord. Surv. Londonderry 1:11 
(1837). 
Callithamnion bipinnatum P. Crouan & H. Crouan, Fl Finistere : 136 (1867). 

See comments on pp. 123; 131. 
Callithamnion caudatum J. Agardh in Linnaea 15 : 46 (1841). 
Ceramium felicii Bonnemaison in Mem. Mus. Hist. nat. Paris 16 : 129 (1828). 
Ceramium tetricum /? pectinatum C. Agardh, Syst. Alg.: 141 (1824). 

3. Rejections from the genus 

In a few cases, examination of type material has disclosed that the entities in question must 
be rejected from the genus Callithamnion. For each, the reason for rejection is stated: 

Callithamnion fallax P. Crouan & H. Crouan 

Type locality: rade de Brest (Crouan & Crouan, 1867 : 135). 

Holotype: CO*, Brest, 27 December 1 856. 

Comments: the holotype consists of depauperate scraps of Antithamnion sp. 

Callithamnion parvulum P. Crouan & H. Crouan 

Type locality: 'Sur le Codium spongiosum . . . rade de Brest' (Crouan & Crouan, 
1867: 135). 

Holotype: CO, Crouan, Brest, 4 March 1858. 

Other material: LD, Herb. Alg. Agardh 1 7973, possibly a duplicate of the holotype. 

Comments: the holotype has the aspect of Spermothamnion, rather than Callithamnion, 
and terminal procarps are present. 

*Abbreviations relating to herbaria are listed in the acknowledgements (pp. 1 38-1 39). 



CALLITH AMNION IN THE BRITISH ISLES 131 

Callithamnion tetricum y minus C. Agardh 

Type locality: Pornic, coll. Desvaux (C. Agardh, 1828 : 180) 
Holotype: LD, Herb. Alg. Agardh 18514. 

Comments: the specimen is of Pleonosporium borreri, as was suggested by J. Agardh 
(1851 :53). 



Typifications 

An attempt has been made to typify every name relevant to the present study. The results are 
listed in alphabetical order, initially by genus and then by species. 

Callithamnion acrospermum J. Agardh 

Type locality: 'in Oceano atlantico ad oras Galliae' (J. Agardh, 1851 : 52). 
Lectotype: LD, Herb, Alg. Agardh 18295. 
Other material: LD, Herb. Alg. Agardh 18296.. 

Callithamnion affme Harvey 

Type locality: Bute, coll. Greville (Harvey in Hooker, 1833 : 344). 

Lectotype specimen: TCD, Greville, Bute, undated, annotated 'Plant of Br. Flora H' in 
Harvey's hand. 
Other material: TCD, three specimens. BM(K), one specimen. 

Callithamnion arachnoideum C. Agardh 

Type locality: 4 in mari Atlantico ad Noveboracum, unde misit Torry' (C. Agardh, 
1828: 181). 

Lectotype: LD, Herb. Alg. Agardh 18435. 

Other material: LD, Herb. Alg. Agardh 18436 [mica preparation, probably of 18435], 
18443. 

Callithamnion arachnoideum Griffiths ex Moore, nom. Meg. 

Type locality: 'near the mouth of the Ban n' (Moore, 1837 : 11). 
Type material: it has not been possible to locate any original material. 

Callithamnion arnottii Trevisan, nom. Meg. 

Comments: C. arnottii (as 4 C. ArnottF) was proposed by Trevisan (1 845 : 77) as a substitute 
name for C. byssoides. It is, therefore, a superfluous and illegitimate name which must be 
typified by the type of C. byssoides (q.v.). 

Callithamnion bipinnatum P. Crouan & H. Crouan 

Type locality: rade de Brest (Crouan & Crouan, 1 867 : 1 36). 
Lectotype: CO, Crouan, rade de Brest, Aug. 1 850. 
Other material: CO, 3 specimens. 

Comments: for discussion of the possible status of this taxon in different parts of Europe, 
see Price ( 1 978) and Rueness & Rueness ( 1 980). 

Callithamnion brodiaei Harvey 

Type localities: Forres, Coll. Brodie; Northumberland, coll. Robertson; Torquay, coll. 
Griffiths & Cutler (Harvey in Hooker, 1833: 340). 



132 P. S. DIXON & J. H. PRICE 

Lectotype: TCD, Brodie, Lossiemouth, undated, annotated as 'authentic' in Harvey's 
hand. 

Other material: TCD, three specimens. LD, Herb. Alg. Agardh 19018, 19025. GL, two 
specimens. 

Comments: the original treatment contains two curious misstatements. The Brodie 
specimens were collected at Lossiemouth according to Brodie's annotations, not at Forres, 
which was where he lived and is a few kilometres from the sea. The Northumberland 
specimen is annotated as having been collected by 'Mr. Robinson', whereas the collector 
was quoted as 'Mr. Robertson' in the published text. 

Callithamnion byssoides Arnott ex Harvey 

Type localities: 'Whitsand bay, G.A.W. Arnott, Esq. Devonshire, Mrs. Griffiths' (Harvey 
in Hooker, 1833:342). 

Lectotype: TCD, Arnott, Whitsand Bay, 1829, annotated 'original plant of Br. Flora 
WHH' (in Harvey's hand). 

Other material: TCD, one specimen. LD, Herb. Alg. Agardh 18407. GL, seven 
specimens, possibly duplicates. 

Callithamnion caudatum J. Agardh 

Type locality: 'in mari mediterraneo ad littora Galloprovinciae; rupibus insulae St. 
Marguerite' (J. Agardh, 1841 : 46). 

Lectotype: LD, Herb. Alg. Agardh 18386. 

Other material: LD, Herb. Alg. Agardh 18390. Numerous other specimens exist (BM, 
TCD, etc.), distributed by J. Agardh, but it would seem best to exclude these because of 
doubts as to their provenance. 

Callithamnion decompositum J. Agardh 

Type locality: 'in oceano atlantico ad oras Galliae' (J. Agardh, 1851: 45). 

Holotype: LD, Herb. Alg. Agardh 18964. 

Comments: this entity is often attributed, partly or completely, to Grateloup, due to a 
misinterpretation of the comment by J. G. Agardh that Grateloup had given the manuscript 
name Mertensia decomposita to the specimen used as the basis for the original treatment. 

Callithamnion fruticulosum J. Agardh 

Type locality: 'in sinu Codano' (J. Agardh, 1841 : 46). 

Lectotype: LD, Alg. Agardh 19047, bearing TYPUS slip attributed to T. Levring. 

Other material: LD, Herb, Alg. Agardh 19046, 19048. TCD, J. Agardh, undated, a mica 
preparation in a packet annotated 'Callith. fruticulosum Ag. MSCR e sinu Codano J. 
Agdh.' in J. Agardh's hand. 

Callithamnion furcellariae J. Agardh 

Type locality: 'in Furcellaria parasiticum ad oras Bahusia' (J. Agardh, 1851 : 37). 

Lectotype: LD, Herb. Alg. Agardh 19087. 

Other material: LD, Herb. Alg. Agardh 19088, 19089. 

Comments: a TYPUS label has been attached to Herb. Alg. Agardh 19085, but it is not 
clear why this was selected because it makes reference to Solenia, not Furcellaria, which is 
cited on the specimen cited above as lectotype. 

Callithamnion grande J. Agardh 

Type locality: 'ad rupes insularum Stoechadum' (J. Agardh, 1842 : 73). 
Holotype: LD, Herb. Alg. Agardh 19277. 



CALLITH AMNION IN THE BRITISH ISLES 133 

Callithamnion grevillei Harvey 

Type localities: 'Firth of Forth, Dr. Greville, Berwick, Dr. Johnston' (Harvey in Hooker, 
1833:345). 

Lectotype: TCD, Greville, no locality, undated, a packet containing a mica preparation, 
annotated 'Plant of Br.Fl.' in Harvey's hand. 

Callithamnion harveyanum J. Agardh 

Type localities: Callithamnion harveyanum was first used by J. Agardh (1841 : 45), who 
claimed that the plant treated by Harvey under the name Callithamnion granulatum was 
distinct from that taxon. As J. Agardh did not provide a formal diagnosis, the original 
description must be that given by Harvey, and the type localities those quoted by the latter, 
viz. Appin [in error, cf. comment belowl, coll. Carmichael; Whitsand Bay, coll. Arnott. 

Lectotype: TCD, Carmichael, StafTa, July 1826, annotated 'Specimen described in British 
Flora as C granulatum is not that plant WHH' in Harvey's hand. 

Other material: LD, Herb. Alg. Agardh 18339. 

Comments: the lectotype and the LD specimen are both annotated to indicate that they 
were collected by Carmichael at StafTa, not Appin, as indicated in the Harvey treatment. 

Callithamnion hiemale Kjellman ex Kylin 

Type locality: Kristineberg (Kylin, 1907 : 170). 
Holotype: LD, Kylin, Kristineberg, 9 December 1905. 

Callithamnion lanosum Harvey 

Type localities: Tor Abbey, Mrs. Griffiths. Killiney, W. H. Harvey' (Harvey in Hooker, 
1833:341). 

Lectotype: TCD, Harvey, Killiney, undated. 

Other material: TCD, four specimens. BM, one specimen. LD, Herb. Alg. Agardh 18257, 
18258. 

Comments: Harris (1966) cited the type locality for this entity as 'Beach below Tor Abbey, 
Menai Straits, N. Wales', a geographically meaningless aggregation of localities that bears no 
relation to the original treatment. 

Callithamnion octosporum C. Agardh 

Type locality: 'Ad litora Galliae maris Atlantici, ut ad Port Louis' (C. Agardh, 
1828: 177). 

Holotype: LD, Herb. Alg. Agardh 18493. 

Other material: LD, Herb. Alg. Agardh 18492, a mica preparation apparently derived 
from 18493. 

Callithamnion polyspermum C. Agardh 

Type locality: 'Ad Finisterre Galliae, unde misit Bonnemaison' (C. Agardh, 1828 : 169). 

Holotype: LD, Herb. Alg. Agardh 18236. 

Other material; LD, Herb. Alg. Agardh 18235, a mica preparation apparently derived 
from 18236. PC, F, QUI, several specimens occur which may possibly represent duplicates 
of the holotype indicated above, since they were received from Bonnemaison and referred 
by him, as was the holotype, to ' Lamourouxia polysperma' (a nomen nudum). 

Callithamnion pseudobyssoides P. Crouan & H. Crouan 

Type locality: 'sur le Plocaria confervoides . . . Kervallon' (Crouan & Crouan, 1 867 : 1 36). 

Lectotype: CO, Crouan, Kervallon, undated. 

Other material: CO, one specimen. LD, Herb. Alg. Agardh 18426. 



134 P. S. DIXON & J. H. PRICE 

Callithamnion roseum /? tenue Lyngbye 

Type locality: Faeroe (Lyngbye, 1819 : 126). 

Lectotype: LD, Herb. Alg. Agardh 18997. 

Other material: LD, Herb. Alg. Agardh 18994. C, Lyngbye, 'Hoyviig prope Thorshavn 
Faeroe', 11 June 1817. PC, Lyngbye, Faeroe, undated, labelled 'Callithamnion roseum 
P tenue', in Lyngbye's hand. 

Comments: Lyngbye and C. Agardh obviously exchanged material extensively; C. Agardh 
made use of exchange material as the basis for both Ceramium scopulorum and 
Callithamnion scopulorum (q.v.). 

Callithamnion scopulorum C. Agardh 

Type locality: Faeroe (C. Agardh, 1828 : 166). 

Lectotype: LD, Herb. Alg. Agardh 18997. 

Other material: LD, Herb. Alg. Agardh 18994. C, Lyngbye, 'Hoyviig prope Thorshavn 
Faeroe', 11 June 1817. PC, Lyngbye, Faeroe, undated, labelled 'Callithamnion roseum f$ 
tenue' m Lyngbye's hand. 

Comments: as discussed under Callithamnion roseum /? tenue (q.v.), C. Agardh made use 
of Lyngbye exchange material of the latter entity as the basis for Callithamnion scopulorum. 
The original treatment of the latter quoted Callithamnion roseum tenue Lyngb.' in its 
synonymy, although there is no mention of Ceramium scopulorum (q.v.), which had already 
been based (C. Agardh, 1 824) on the same material. 

Callithamnion spiniferum Kylin 

Type localities: Hogardsgrund, Varberg, Morup (Kylin, 1907 : 159). 
Lectotype: LD, Kylin, Morup, 21 February 1903. 
Other material: LD, Kylin, Hogardsgrund, 6 July 1907. 

Callithamnion spinosum Harvey 

Type locality: Sidmouth, coll. Griffiths (Harvey in Hooker, 1833 : 345). 

Holotype: TCD, Griffiths, Sidmouth, undated, annotated 'Cal. spinosum H. in Br. Flora. 
Sent by Mrs. Griffiths to Dr. Greville from Sidmouth — my only specimen' in Harvey's hand. 

Comments: Harris (1966) cited as holotype a specimen at TCD, undefined as to locality 
and lacking further identifying data. That specimen is of no importance to the correct 
typification. 

Callithamnion spongiosum Harvey 

Type localities: Torquay, coll. Griffiths; Kingston, coll. Harvey; Killiney, coll. Harvey 
(Harvey in Hooker, 1833 : 346). 

Lectotype: TCD, Griffiths, Meadfoot (Torquay), June 1832. 

Other material: TCD, five specimens. BM, two specimens. BM(K), one specimen. GL, one 
specimen. LD, Herb. Alg. Agardh 19249. 

Callithamnion tetragonum fl firmius C. Agardh 

Type locality: Trishwater [sic] Bay' (C. Agardh, 1828 : 176). 
Holotype: LD, Herb. Alg. Agardh 18341. 

Other material: LD, Herb. Alg. Agardh 18342, a mica preparation apparently prepared 
from 18341. 

Callithamnion tripinnatum C. Agardh 

Type locality: 'Mertensia tripinnata Gratel. Msct. In Mari Atlantico, unde specimen misit 
Grateloup' (C. Agardh, 1828 : 168). 



CALLITH AMNION IN THE BRITISH ISLES 135 

Holotype: LD, Herb. Alg. Agardh 18975. 

Comments: citation of the authority is confused. Direct attribution to Grateloup is 
incorrect as his name is a nomen nudum, while citation in the form 'Gratel. ex. C. Ag.' is also 
incorrect in that C. Agardh took up only the epithet and not the whole binomial from 
Grateloup (see International Code of Botanical Nomenclature, Recommendation 46C. 1). 

Ceramium brachiatum Bonnemaison, nom. illeg. 

Comments: the original treatment (Bonnemaison, 1828:136) made reference to 
collections from three localities on the Atlantic or Channel coasts of France, to another from 
the Mediterranean, and to two specimens collected in the Canary Islands and at the Cape of 
Good Hope. More significantly, it placed in synonymy binomials which lead back to 
Conferva tetragona Withering and made no mention of the latter elsewhere in the text, so 
that these synonyms must be taken as references to type. There is no option but to dismiss 
Ceramium brachiatum as a superfluous and illegitimate name which must be typified by the 
type of Conferva tetragona. 

Ceramium congestum Bonnemaison, nom. illeg. 

Comments: the original treatment (Bonnemaison, 1828 : 125) made reference to 
collections from nine localities on the Channel and Atlantic coasts of France. The critical 
feature of the treatment is that it placed in the synonymy Conferva tetrica Dillwyn (1806). 
On the basis of that citation, one must dismiss Ceramium congestum as a superfluous and 
illegitimate name, to be typified by the type of Conferva tetrica. 

Ceramium dudresnayi Bonnemaison, nom. illeg. 

Comments: the original treatment (Bonnemaison, 1828 : 125) made reference to various 
specimens but also to several previously-published binomials, of which the oldest is 
Conferva purpurascens Hudson. Therefore Ceramium dudresnayi is a superfluous and 
illegitimate name which must be typified by the type of Conferva purpurascens. 

Ceramium felicii Bonnemaison 

Type locality: 'Dieppe' (Bonnemaison, 1828 : 129). 

Lectotype: PC, Gaillon, Dieppe, undated, annotated as 'Ceramium felicii Gaillon mnscr' 
in Bonnemaison's hand. 

Other material: specimens were distributed widely by Gaillon, although he never 
provided a description. The greatest number were issued as specimen 203 of Desmaziere's 
"Plantes Cryptogames" but others, not from this source, occur at CO (ex Crouan) and at LD 
(Herb. Alg. Agardh 19016, also ex Crouan). 

Ceramium fruticulosum Roth 

Type locality: not stated (Roth, 1800 : 183), but as the original material was collected by 
Wulfen it was probably from the Adriatic. 

Type: typification is not currently possible as there is no evidence at either B or W to 
indicate that the original material is still in existence and no illustration is provided. 

Comments: Roth (1800) compared his alga with Conferva marina nodosa of Dillenius 
(1741), the type of Conferva purpurascens Huds., and in a later treatment (Roth, 1806 : 147) 
actually cited the latter in his treatment of Ceramium fruticulosum. Wulfen (1803) 
independently described Conferva fruticulosa (q.v.), and it is probable that the two entities 
are based on fragments of the same original gathering. 

Ceramium granulatum Ducluzeau 

Type locality: Sette (Ducluzeau, 1806 : 72). 



136 P. S. DIXON & J. H. PRICE 

Type: typification is not currently possible as nothing is known of the location of any 
Ducluzeau material and no illustration, although announced, is provided with the original 
treatment. 

Ceramium guttatum Bonnemaison 

Type localities: Penmark, Cherbourg (Bonnemaison, 1828 : 137). 

Lectotype: PC, locality not indicated, Bonnemaison, undated, labelled 'Ceramium 
guttatum B n sp. nov.', in Bonnemaison's hand, and filed under Callithamnion tetragonum. 

Ceramium roseum Roth 

Type locality: Bayonne (Roth, 1798 : 47). 

Type: indisputable type material not located. Neotype designated below. 

Comments: the existence of Roth's herbarium is at present a matter of doubt. Usually, it is 
assumed to have been lost in the bombing of the Berlin-Dahlem Museum (Pilger, 1953); this 
seems true of specimens of Ceramium roseum. There is a specimen at LD (Herb. Alg. 
Agardh 18466), labelled 'Ceramium roseum Roth Bayonne dedit Roth ipse non rosea Dillw' 
in C. Agardh's hand, but the status of this is uncertain. Subsequent interpretations (Dillwyn, 
1802; Smith, 1802) were based on a specimen collected in 1797 by Sowerby at Yarmouth 
and sent to Roth for comment by Dawson Turner. This specimen, which Roth identified as 
'Ceramium roseum\ passed to K (now at BM), with a duplicate of the original Sowerby 
collection at BM. Although not part of the type collection, these specimens help to unravel 
subsequent interpretations of this taxon, of which there have been several. This K(BM) 
specimen is here designated as of neotype status. 

Ceramium scopulorum C. Agardh 

Type locality: Faeroe (C. Agardh, 1824 : 132). 

Lectotype: LD, Herb. Alg. Agardh 18997. 

Other material: C, Lyngbye, 'Hoyviig prope Thorshavn Faeroe', 11 June 1817. PC, 
Lyngbye, Faeroe, undated, labelled 'Callithamnion roseum tenue" in Lyngbye's hand. 

Comments: as discussed under Callithamnion roseum /? tenue (q.v.), Lyngbye and C. 
Agardh exchanged material and C. Agardh made use of material of the latter entity as the 
basis for Ceramium scopulorum. The original treatment of the latter quoted 'Callithamnion 
roseum tenue, Lyngb.' in its synonymy. The same material was later used by C. Agardh as 
the original material for Callithamnion scopulorum (q.v.), although the latter description 
makes no reference to the earlier Ceramium scopulorum. 

Ceramium tenuissimum Bonnemaison 

Type localities: St-Pol-de-Leon, Brest, St. Malo (Bonnemaison, 1828 : 132). 

Provisional lectotype: PC, Bonnemaison, no locality, undated, annotated 'Ceramium 
tenuissimum B n ' in Bonnemaison's hand, filed as Seirospora tenuissimum. 

Comments: despite widespread search at QUI, it was not possible to locate any authentic 
material in what remains of Bonnemaison's herbarium. Presumably the specimen cited 
above as provisional lectotype is that referred to by Harris (1966). 



Ceramium tetricum /? pectinatum C. Agardh 

Type locality: 'Ad orasGalliae, Africae Borealis'(C. Agardh, 1824 : 141). 
Lectotype: LD, Herb. Alg. Agardh 18307. 
Other material: LD, Herb. Alg. Agardh 1 8309. 



CALLITH AMNION IN THE BRITISH ISLES 1 37 

Ceramium versicolor C. Agardh nom. Meg. 

Comments: the original treatment (C. Agardh, 1824:140) cites three previously 
published binomials, one identification, and the manuscript source of the epithet. Despite 
widespread search, we have been unable to locate the type material of any of the previous 
binomials. Ceramium versicolor is a superfluous and illegitimate name, incapable of typifi- 
cation, which must be dismissed from consideration. 

Conferva arbuscula Dillwyn 

Type localities: Ballycastle, coll. Brown; Bantry Bay, coll. Hutchins (Dillwyn, 1807, pi. 
85). 

Lectotype: BM(K), ? Brown, North coast of Ireland, undated. 

Other material: BM, 2 specimens. GL, 2 specimens. 

Comments: the typification has been discussed in detail by Dixon (1960, 1964). The 
choice of lectotype was made on the basis of the statement by Dillwyn (1809 : 35) that 
Dawson Turner provided the original description of the entity. The position became 
confused as a consequence of a later figure (Dillwyn, 1809, pi. 6G) based on a specimen of 
Dasya, so that the epithet arbuscula came to be applied in both Callithamnion and Dasya. 
Despite previous resolution of this problem (Dixon, 1960, 1964), Harris (1966) stated that he 
had been unable to locate original material. 

Conferva corymbosa Smith 

Type localities: Brighthelmstone [=Brighton], coll. Borrer; Kingsbridge, coll. Gibbs; 
Bantry Bay, coll. Hutchins (Smith, 181 1, pi. 2352). 

Lectotype: BM, Borrer, Brighton, July 1811. 

Comments: Harris (1966) quoted the type locality as Kingsbridge and cited as holotype a 
specimen 'in the British Museum of Natural History, no. 271'. The original treatment is 
based on three collections, so that holotype status for any specimen is impossible. It would 
appear that this author was misled by the seven irrelevant specimens, in the type folder at 
BM, mounted on the same sheet as the Borrer specimen. The specimen cited by Harris is an 
example from "Algarum Fasciculi' distributed by J. Cocks, and has no relevance whatsoever 
to the typification of Conferva corymbosa. 

Conferva fruticulosa Wulfen 

Type locality: Tnsulae Gradensis' (Wulfen, 1803 : 26). 

Type: typification is not currently possible as the original material cannot be located, and 
no illustration is provided with the original treatment. 

Comments: Wulfen stated that his material was similar to the illustration of Conferva 
marina nodosa of Dillenius (1742). Roth (1800) described Ceramium fruticulosum on the 
basis of material supplied by Wulfen and also commented on the relationship to the same 
Dillenian entity. Conferva fruticulosa Wulfen and Ceramium fruticulosum Roth are 
independent names, based on different types, although these are probably fragments of the 
same original gathering. 

Conferva hookeri Dillwyn 

Type localities: Cawsie, coll. Hooker and Borrer; Holyhead, coll. Davies; Bantry Bay, coll. 
Hutchins (Dillwyn, 1809 : pi. 106). 

Lectotype: TCD, Hooker and Borrer, Cawsie, undated, labelled 'Br. Flora C. Hookeri 
Cawsie Hook. Borrer', in Harvey's hand. 

Other material: BM(K), three specimens. 

Comments: Harris (1966) cited a K specimen, supposedly from Holyhead, Anglesey. 
None of the K material mentioned above is so annotated and the specimen at TCD is the 
legitimate choice of lectotype. 



138 P. S. DIXON & J. H. PRICE 

Conferva purpurascens Hudson 

Type locality: unspecified (Hudson, 1778 : 600). 

Holotype: OXF, Dillenian material of Conferva marina nodosa. 

Comments: Hudson's description is based exclusively on the plant described previously by 
Dillenius (1742) as Conferva marina nodosa . . . Harris (1966) came to a similar conclusion, 
although he considered the Dillenian material as of neotype status. Although Hudson cited 
no precise locality in his treatment of Conferva purpurascens, Dillenius (1742 : 36) indicates 
that his material was collected near 'Lhanfaethly', Anglesey, which is therefore the type 
locality for the Hudson taxon. 

Conferva seposita Gunnerus 

Type localities: 'ad Hoburgam Gotlandiae; in man sundmorico [near Alesund, Norway]; 
prope Lhanfaethly in Mona Insula [Anglesey]' (Gunnerus, 1772 : 116). 

Lectotype: OXF, Green, Lhanfaethly, undated, in Dillenius's Historia Muscorum 
herbarium. 

Comments: the lectotype selected is also the lectotype of Conferva purpurascens Hudson 
(q.v.). 

Conferva tetragona Withering 

Type locality: 'found by Major Velley and Mr. Stackhouse at the Bill of Portland' 
(Withering, 1796:405). 
Provisional lectotype: GL, Velley, Isle of Portland, undated. 

Conferva tetrica Dillwyn 

Type locality: 'Common about the Mumbles and in other parts of the Peninsula of Cower' 
(Dillwyn, 1806, pi. 81). 

Lectotype: WELT, Herb. Silvanus P. Thompson A 4502, Dillwyn, Swansea, 1806. 

Ellisius glaber S. F. Gray, nom. Meg. 

Comments: E. glaber is a superfluous and illegitimate name, in that Conferva arbuscula 
Dillw. is cited as a synonym (Gray, 1821 : 333). The genus Ellisius was established with two 
species, E. glaber and E. coccineus, the latter based on Conferva coccinea Hudson 
[= Heterosiphonia plumosa (Ellis) Batters]. Heterosiphonia Montagne (1842) has been 
conserved against Ellisius. 

Phlebothamnion faroense Kutzing 

Type locality: 'Faeroe' (Kutzing, 1864 : 30). 

Holotype: L, Herb. Lugd. Bat. 940.237.1 16, Lyngbye, Faeroe, undated. 

Phlebothamnion scoticum Kutzing 

Type locality: Aberdeen (Kutzing, 1864 : 30). 

Holotype: L, Herb. Lugd. Bat. 940.237. 118, Dickie, Aberdeen, June 1 843. 



Acknowledgements 

A work of this magnitude could not have been conducted without the active support and co-operation 
of many institutions and collections. We wish to express our indebtedness to the Directors and 
Curators of the following institutions: School of Plant Biology, University College of North Wales, 
Bangor (UCNW); Laboratoire de Biologie Marine, Concarneau (CO); Botanical Museum and 
Herbarium, University of Copenhagen (C); University College, Cork; School of Botany, Trinity 



CALLITH AMNION IN THE BRITISH ISLES 1 39 

College, Dublin (TCD); Herbarium, Royal Botanic Garden, Edinburgh (E); Istituto Botanico, Firenze 
(FL); Department of Botany, University of Glasgow, Glasgow (GL); Royal Botanic Gardens, Kew (K) 
[algal material now at BM]; Rijksherbarium, Leiden (L); Merseyside County Museum, Liverpool (LIV); 
British Museum (Natural History), London (BM); Botanical Museum, Lund (LD); The Manchester 
Museum, The University of Manchester, Manchester (MANCH); Hancock Museum, Newcastle-upon- 
Tyne (HAMU); Botanical Museum, Oslo (O); Fielding-Druce Herbarium, Oxford (OXF); Laboratoire 
de Cryptogamie, Museum National d'Histoire Naturelle, Paris (PC); Eastgate House Museum, 
Rochester; Station Biologique, Roscoff; Sunderland Museum, Sunderland; County Museum, Truro; 
Institute of Systematic Botany, University of Uppsala, Uppsala (UPS); Museo Civico di Storia 
Naturale, Venezia; National Museum, Wellington (WELT); Naturhistorisches Museum, Wien (W). 

Financial support has been received from various sources over the past 20 years, including the Joint 
Committee on Research of the University of Liverpool; the Browne Fund of the Royal Society, London; 
the British Council; the Faculty Research and Travel Fund of the University of California; the 
National Science Foundation; and the British Museum (Natural History). 

Many individuals have assisted with discussions and with the provision of information and 
specimens. We would like to thank P. Edwards, W. F. Farnham, M. D. Guiry, J. D. Guiterman, 
C. E. L. Hepton, D. E. G. & L. M. Irvine, D. M. John, T. Levring, C. Maggs, O. Morton, W. Prud' 
homme van Reine, J. and M. Rueness, P. C. Silva, I. Tittley, J. A. West & A. Whittick for this 
assistance. 



References 

Agardh, C. A. 1 824. Systema Algarum. xxxviii + 3 12 pp. Lund. 

1 828. Species Algarum rite cognitae ... 2 (1), lxxvi + 1 89 pp. Gryphiae. 

Agardh, J. G. 1841. In historiam algarum symbolae. Linnaea 15 : 1-50; 443-457. 

1 842. Algae Maris Mediterranei et Adriatici ... x + 1 64 pp. Paris. 

1851. Species Genera et Ordines Algarum ... 2 (1) Algae Florideae complectens. xi + 336 + 

[addenda 337-35 1 pp.] + index. Lundae. 

1892. Analecta Algologica. Acta Univ. lund. 28 (6) : 1-182. 



Boddeke, R. 1958. The genus Callithamnion Lyngb. in the Netherlands. A taxonomic and oecological 

study. Acta bot. need. 7 : 589-604. 
Bonnemaison, T. 1828. Essai sur les Hydrophytes loculees (ou articulees) de la famille des Epidermees 

et des Ceramiees. Mem. Mus. Hist. nat. Paris 16 : 49-1 58. 
Chadefaud, M. 1954. Sur la morphologie de quelques Ceramiacees. Revue algol. II, 1 : 71-87. 
Chadefaud, M. 1979. L'evolution de la structure cladomienne chez les Charales et les Ceramiales. 

Etude comparative. Revue algol. II, 14 : 253-273. 
Cotton, A. D. 1912. 15 Marine Algae. In: R. L. Praeger, A biological survey of Clare Island in the 

county of Mayo, Ireland and the adjoining district. Proc. R. Ir. A cad. 31, sect 1 (15): 1-178. 
Crouan, P. L. & Crouan, H. M. 1 867. Florule du Finistere. x + 262 pp. Paris & Brest. 
Dillenius, J. J. 1 742 [' 1 74 1']. Historia muscorum ... [2] + 1-2 + iii-xvi + 576 pp. Oxford. 
Dillwyn, L. W. 1802-1809. British Confervae; or Coloured Figures and Descriptions of the British 

Plants referred by Botanists to the Genus Conferva. [2] + 87 + [23 1] pp., pis 1-109, A-G. London. 
Dixon, P. S. 1960. Taxonomic and nomenclatural notes on the Florideae II. Bot. Notiser 

113:295-319. 

1964. Taxonomic and nomenclatural notes on the Florideae. IV. Bot. Notiser HI : 56-78. 

1971. Cell enlargement in relation to the development of thallus form in Florideophyceae. Br. 

phycol.J.6: 195-205. 
Drew, K. M. 1939. An investigation of Plumaria elegans (Bonnem.) Schmitz with special reference to 

triploid plants bearing parasporangia. Ann. Bot. II, 1 : 463-476. 
Ducluzeau, J. A. P. 1806 ['1805']. Essai sur I'histoire naturelle des Conferves des environs de 

Montpellier. 89 pp. Montpellier. 
Edwards, P. 1969. The life history of Callithamnion byssoides in culture. J. Phycol. 5 : 266-268. 
1979. A cultural assessment of the distribution of Callithamnion hookeri (Dillw.) S. F. Gray 

(Rhodophyta, Ceramiales) in nature. Phycologia 18 : 251-263. 
Feldmann-Mazoyer, G. 1941 ['1940']. Recherches sur les Ceramiacees de la Mediterranee occidentale. 

510 pp. Alger. 
Foldvik, N. 1963. En undersokelse over okologi og livs-syklus hos gruppen Callithamnion i omrddet 

omkring Biologisk stasjon, Espegrend. Cand. real, thesis, University of Oslo, Norway. 



140 P. S. DIXON & J. H. PRICE 

Gray, S. F. 1821. A Natural Arrangement of British Plants, ... 1. xxviii + 824 pp. London. 

Guiry, M. D. 1978. The importance of sporangia in the classification of the Florideophyceae. In 

D. E. G. Irvine & J. H. Price, Modern Approaches to the Taxonomy of Red and Brown Algae: 

1 1 1-144. London, New York, San Francisco. 
Gunnerus, J. E. 1 772. Flora Norvegica . . . 2 : [2] + 148 + [84] pp. Hafniae. 
Harris, R. E. 1959. A taxonomic study of the genus Callithamnion Lyngbye emend. Ndgeli. Ph.D. 

thesis, University of Manchester. 

1962. Contribution to the taxonomy of Callithamnion Lyngbye emend. Naegeli. Bot. Notiser 

115: 18-28. 

1966. Contribution to the genus Callithamnion Lyngbye emend. Naegeli: taxonomy of the species 

indigenous to the British Isles. Advg Front. PL Sci. 14: 109-131. 
Harvey, W. H. 1 833. Div. II Confervoideae. Div. Ill Gloiocladeae. In W. J. Hooker, The English Flora 

of Sir James Edward Smith 5. x + 430 pp. London. 
— - 1 847 [' 1 846-5 1 *\. Phycologia Britannica . . . 1 . xv + viii pp. + 1 20 pis (with text). London. 
Hassinger-Huizinga, H. 1952. Generationswechsel und Geschlectsbestimmung bei Callithamnion 

corvmbosum (Sm.) Lyngb. Arch. Protistenk. 98 : 91-124. 
Hudson, W. 1778. Flora Anglica; ... Ed. Altera, 1. [4] + xxxviii + [1] + 396 pp; 2. 397-690 pp. London. 
Kapraun, D. F. 1978. Field and culture studies on growth and reproduction of Callithamnion byssoides 

(Rhodophyta, Ceramiales) in North Carolina. J. Phycol. 14 : 21-24. 
Kiitzing, F. T. 1 843. Phycologia generalis . . . xvi + 1-144 + xvii-xxxii + 145-1 58 + [1] pp. Leipzig. 

1 849. Species Algarum. vi + 922 pp. Lipsiae. 

1864. Tabulae Phycologicae ... 14. i + 35 pp., 100 pis. Nordhausen. 

Kylin, H. 1907. Studien iiber die Algenflora der schwedischen Westkilste. iv + 287 + [2] pp. Upsala. 

1956. Die Gattungen der Rhodophyceen. xv + 673 pp. Lund. 

L'Hardy-Halos, M.-Th. 1970. Recherches sur les Ceramiacees (Rhodophycees-Ceramiales) et leur 

morphogenese. I. — Structure de l'appareil vegetatif et des organes reproducteurs. Revue gen. Bot. 

77:211-287. 

1971a. Recherches sur les Ceramiacees (Rhodophycees — Ceramiales) et leur morphogenese. II. 

Les modalites de la croissance et les remaniements cellulaires. Revue gen. Bot. 78 : 201-256. 

1971b. Recherches sur les Ceramiacees (Rhodophycees-Ceramiales) et leur morphogenese. III. — 



Observations et recherches experimentales sur la polarite cellulaire et pour la hierarchisation des 
elements de la fronde. Revue gen. Bot. 78 : 407-49 1 . 

1971c. Recherches sur les Ceramiacees (Rhodophycees, Ceramiales) et sur quelques aspects de 



leur morphogenese (1). Bull. Soc. sci. Bretagne 46 : 99-1 12. 
Lyngbye, H. C. 1819. Tentamen hydrophytologicae danicae . . . xxxii + 248 pp. Hafniae. 
Mathias, W. T. 1927. The cytology of Callithamnion. Rep. Br. Ass. Advmt Sci. 1927 (95) : 380. 

1928. The cytology of Callithamnion brachiatum Bonnem. Pubis Hartley bot. Labs Lpool Univ. 

5 : 1-27. 

1932. The cytology of Callithamnion brachiatum Bonnem. Ann. Bot. 46 : 185-187. 

1935. The life-history and cytology of Phloeospora brachiata Born. Pubis Hartley bot. Labs Lpool 

Univ. 13 : 1-28. 
Miranda, F. 1934. Materiales para una flora marina de las rias bajas gallegas. Boln. R. Soc. esp. Hist. 

nat.34: 165-180. 
Montagne, C. 1842. Prodromus Generum Specierumque Phycearum novarum, in Itinere ad Polum 

Antarcticum ... 16 pp. Parisiis. 
[Moore, D.[ 1837. Botany. In Anon., Ordnance Survey Notices: 6-13. In T. A. Larcom, Ordnance 

Survey of the County of Londonderry . . . 1 : 1-16. Dublin. 
Murray, S. N. & Dixon, P. S. 1972. The effect of light intensity and light period on the development of 

thallus form in the marine red alga Pleonosporium squarrulosum (Harvey) Abbott (Rhodophyta: 

Ceramiales). I. Apical cell division — main axes. J. exp. mar. Biol. Ecol. 13 : 1 5-27. 
& 1975. The effects of light intensity and light period on the development of thallus form in 

the marine red alga Pleonosporium squarrulosum (Harvey) Abbott (Rhodophyta: Ceramiales). II. 

Cell enlargement. J. exp. mar. Biol. Ecol. 19 : 165-176. 
Nageli, C. 1847. Die neuern Algensysteme und versuch zur Begriindung eines eigenen Systems der 

Algen und Florideen. Neue Denkschr. Allg. schweiz. Ges. ges. Naturw. 9 (unnumbered art no. 2) 

275 pp. 

1862 ['1861']. Beitrage zur Morphologie und Systematik der Ceramiaceen. Sber. baver. Akad. 

Wi.s-.s-. 1861 (2) -.297-415. 

Newton, L. 1 93 1 . /I Handbook of the British Seaweeds, xiii + 478 pp. London. 



CALLITHAMNION W THE BRITISH ISLES 141 

Pilger, R. 1953. Bericht iiber den Botanischen Garten und das Botanische Museum Berlin-Dahlem 

vom 1. Marz 1943 bis 31. Marz 1947. Willdenowia 1:1-21. 
Price, J. H. 1978. Ecological Determination of Adult Form in Callithamnion: Its Taxonomic 

Implications. In D. E. G. Irvine & J. H. Price, Modern Approaches to the Taxonomy of Red and 

Brown Algae: 263-300. London, New York, San Francisco. 
& Tittley, I. 1 978. Marine algae (excluding diatoms). In A. C. Jermy & J. A. Crabbe, The Island of 

Mull a survey of its flora and environment: 19.1-19.37. London. 
Rosenvinge, L. K. 1924. The marine algae of Denmark. Contributions to their natural history. Part III. 

Rhodophyceae III. (Ceramiales). K. danske Vidensk. Selsk. Skr., 1 Raekke, Nat. og Mat. afd. 

7 (3): 285-488. 
Roth, A. W. 1798. Novae Plantarum Species. Archiv Bot. (Romer) 1 (3) : 37-52. 

1800. Catalecta Botanica ... 2 : [10] + 258 + [12] pp. Lipsiae. 

• 1806. Catalecta Botanica . . . 3 : [8] + 350 + [9] pp. Lipsiae. 

Rueness, J. & Rueness, M. 1980. Culture and field observations on Callithamnion bipinnatum and C 

byssoides (Rhodophyta, Ceramiales) from Norway. Sarsia 65 : 29-34. 
Rueness, M. & Rueness, J. 1978. A parasporangium-bearing strain of Callithamnion hookeri 

(Rhodophyceae, Ceramiales) in culture. Norw. J. Bot. 25 : 201-205. 
Smith, J. E. 1802. See Smith &Sowerby, 1790-1814. 

1811. See Smith &Sowerby, 1790-1814. 

& Sowerby, J. 1 790-1 8 14. English Botany; ... 36 vols with indices. London. 

Trevisan [di San Leon], V. B. A. 1845. Nomenclator Algarum ... 1. 80 pp. Padoue. 

Westbrook, M. A. 1927. Callithamnion scopulorum C. Ag. J. Bot., Lond. 65 : 129-138. 

1930a. Structure of the nucleus in Callithamnion spp. Feulgens 'Nuclealfarbung' for chromatin. 

Ann.Bot.44: 1012-1015. 

19306. Callithamnion tetricum (Dillw.) Ag. J. Bot., Lond. 68 : 193-203. 

1935. Observations on nuclear structure in the Florideae. Beih. bot. Zbl. 53 (Abt. A) : 564-585. 



Whittick, A. 1973. The Taxonomy, Life History and Ecology of Some Species of the Ceramiaceae 
(Rhodophyta) in the North-west Atlantic. 368 pp. Ph.D. thesis, Department of Biology, Memorial 
University of Newfoundland. 

1978. The life history and phenology of Callithamnion corymbosum (Rhodophyta: Ceramiaceae) 

in Newfoundland. Canad. J. Bot. 56 : 2497-2499. 

& West, J. A. 1979. The life history of a monoecious species of Callithamnion (Rhodophyta, 



Ceramiaceae) in culture. Phycologia 18 : 30-37. 
Withering, W. 1796. An Arrangement of British Plants. 3rd ed. 4: [4] + 418 pp. Birmingham & 

London. 
Wulfen, [F.] X. de 1 803. Cryptogama aquatica. [2] + 64 pp. Lipsiae. 



Addendum 

Since this paper went to press the following references have come to the authors' attention. 

Hoek, C. van den (in press). Distribution groups of benthic marine algae in relation to the temperature 

regulation of their life histories. In J. H. Price, R. W. Sims & P. E. S. Whalley, Evolution, Time and 

Space : the Emergence of the Biosphere. London. 
Price, J. H. (in press). Callithamnion Lyngbye 1819. In G. W. Lawson & D. M. John, The marine algae 

and coastal environment of tropical West Africa. Beih. nov. Hedwigia 70. 
South, G. R. & Hooper, R. G. 1980. A catalogue and atlas of the benthic marine algae of the island of 

Newfoundland. Mem. Univ. Newfoundland occ. Pap. Biol. 3 : 1-136. 



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Parmelia subgenus Amphigymnia 
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GENERAL * 

Parmelia subgenus Amphigymnia (lichens) in East 
Africa 






Hildur Krog 

Botanical museum, Trondheimsveien 23 B, Oslo 5, Norway 
T. D. V. Swinscow 

24 Monmouth Street, Topsham, Exeter, England 

Contents 

Page 

Synopsis 143 

Introduction 144 

Previous collections and research 144 

Taxonomic review 144 

Materials and methods 145 

Results 145 

Morphology 145 

Chemistry 148 

Ecology and distribution 150 

Discussion 153 

Circumscription of the species 153 

Generic concept and infrageneric classification 158 

The East African Amphigymnia species 159 

Key to the species 159 

Descriptions of the species 165 

Acknowledgements 227 

References 227 

Index 229 



Synopsis 

Seventy species of Parmelia subgenus Amphigymnia (Vainio) Dodge are reported from East Africa, 
their ecology and distribution discussed, and a key and descriptions to the species are provided. The 
following new species are described: Parmelia aprica, P. durumae, P. jacarandicola, P. kwalensis, P. 
leonis, P. pardii, P. pigmentifera, P. rava, P. taitae, P. tsavoensis, P. umbrosa, and P. vivida. The new 
name Parmelia indoafra is proposed for Parmotrema indicum Hale, non Parmelia indica Sprengel. 
The following taxa are reduced to synonymy: Parmelia aberrans (Vainio) des Abb., P. 
madagascariacea (Hue) des Abb., and P. nyasensis Dodge with P. xanthina (Mull. Arg.) Vainio; 
Parmelia affluens Hale with P. dilatata Vainio; Parmelia balensis Winnem, P. neolobulascens 
Winnem, and Parmotrema subcompositum Hale with Parmelia subschimperi Hale; Parmelia 
breviciliata Hale and P. procera Steiner & Zahlbr. with P. maclayana Mull. Arg.; Parmotrema 
conferendum Hale and Parmelia leucosemotheta Hue with P. subsumpta Nyl.; Parmelia composita 
Hale and P. spilota Hale with P. euneta Stirton; Parmelia deflectens Kurok. with P. subarnoldii des 
Abb.; Parmelia diversa Hale with P. nilgherrensis Nyl.; Parmelia modesta Hue with P. andina Mull. 
Arg.; Parmelia mordenii Hale with P. praesorediosa Nyl.; Parmelia natalensis Steiner & Zahlbr. and P. 
pseudonilgherrensis Asah. with P. lobulascens Steiner; Parmelia pachyspora Hale with P. amaniensis 
Steiner & Zahlbr.; Parmelia paradoxa Hale with P. inexpectata des Abb.; Parmelia pseudoreticulata 
Tavares and P. simulans Hale with P. reticulata Taylor; Parmelia pseudotinctorum des Abb. with P. 
tinctorum Nyl.; and Parmelia virens Mull. Arg. with P. erubescens Stirton. Parmelia consors Nyl. and 
P. pilosa Stizenb., previously classified in genus Parmelina Hale, are here included in subgenus 



Bull. Br. Mus. nat. Hist. (Bot.) 9 (3) : 143-23 1 



Issued 26 November 1 98 1 



143 



144 H. KROG & T. D. V. SWINSCOW 

Amphigymnia (genus Parmotrema Massal.). The generic concept and infrageneric classification are 
briefly discussed. 



Introduction 

Previous collections and research 

Species of Parmelia subgenus Amphigymnia (Vainio) Dodge form one of the major 
dominant groups of lichens in East Africa. Being large and conspicuous they were frequently 
collected by both botanists and non-botanists on various expeditions during the 19th 
century. New species and infraspecific taxa now classified in Amphigymnia were 
described by contemporary lichenologists (names in parentheses) from material brought 
back to Europe by the following collectors: Hannington (Muller, 1890), Hildebrandt 
(Krempelhuber, 1877, Muller, 1884, 1885), Hoist (Muller, 1894), Liechtenstein & 
Pospischill (Steiner, 1897), Maclay (Muller, 1891), Meyer (Stein, 1888; Muller, 1890), 
Schimper (Muller, 1892), Scott Elliot (Vainio, 1898), Stuhlmann (Muller, 1893, 1894), 
Thomson (Stirton, 1877-78), and Volkens (Muller, 1894). Most of these new taxa, together 
with records of other lichens occurring in our area, were included in an account by 
Stizenberger(1890, 1891, 1893, 1895) of lichens known from Africa at that time. 

In this century Jatta (1909) published an account of lichens from the Ruwenzori 
mountains, and Hue (1916) reported on lichens collected in Kenya by Vicomte de Poncins. 
Steiner & Zahlbruckner (in Zahlbruckner, 1926), Zahlbruckner (1932), and Zahlbruckner & 
Hauman (1936) published on various collections of lichens from the African continent. 
Several of them are pertinent to our area, especially those collected by Brunnthaler, as well 
as those by Fincke, Schroder, Tobler, and F. & R. Wettstein. Cengia Sambo (1937a, b, 1938, 
1940) reported on lichens mainly from Ethiopia, but also from Kenya and Tanzania, and 
Klement (1962) published an account of a small collection of lichens from Tanzania made 
by Drs E. and R. Schuz. 

Dodge (1953, 1959) studied those East African Parmelias lodged in the herbaria of EA, 
FH, and K (now in BM), describing a number of new species. His accounts include many 
species in Amphigymnia which are now accommodated elsewhere. Hale (1965) published a 
world monograph of the subgenus, reducing numerous taxa to synonymy and describing 
many new species. Nearly 30 of the species he treated were then known to occur in East 
Africa. In subsequent publications Hale (1972, 1973, 1977) described several additional 
species from our area. Winnem (1975) treated the 31 species of the subgenus Amphigymnia 
occurring in Ethiopia. A further two species have previously been discussed by the present 
authors (Krog, 1 974; Krog & Swinscow, 1 975). It is noteworthy that most of the authors cited 
were not familiar with the African lichen flora in the field. The main exception is Winnem 
(1975), who based her study of Ethiopian lichens on her own collections and a first-hand 
knowledge of the physiography of the country. 

The aim of the present study is to give a comprehensive account of the species belonging to 
Parmelia subgenus Amphigymnia in East Africa. Special emphasis is placed on the circum- 
scription of the species in the light of characters of the reproductive organs, new chemical 
data, ecological observations, and a knowledge of the variation within local populations. 

Taxonomic review 

Massalongo (1860) segregated the genus Parmotrema Massal., based on Parmelia perforata 
(Jacq.) Ach., from Parmelia Ach. Vainio (1890) introduced Parmelia subgenus Euparmelia 
sect. Amphigymnia Vainio with 1 1 species, but did not designate a type. Dodge (1959) raised 
the section to subgeneric level, noting that a type had not been designated. Hale (1965) 
treated the subgenus monographically and selected Parmelia tinctorum Nyl. as the lectotype 
for it. Hale ( 1 974/?) reintroduced genus Parmotrema Massal., including in it not only most of 



PARMELIA (AMPH/GYMNIA) IN EAST AFRICA 145 

the species which had previously been accommodated in subgenus Amphigymnia (Vainio) 
Dodge, but also the P. reticulata group, previously included in subgenus Parmelia. 



Materials and methods 

The following account is based mainly on collections made by the authors as follows: 
Uganda (T. D. V. S. 1969, 1970, 1971), southern Ethiopia (H. K. 1972), Kenya (H. K. & T. 
D. V. S. 1972, 1973, 1974, H. K. 1976, T. D. V. S. 1 977), and northern Tanzania (H. K. & T. 
D. V. S. 1974). In addition to specimens in institutional herbaria we have been permitted to 
examine material collected in recent years by the following botanists: Lye (Uganda), Moberg 
(Kenya, Tanzania), Pocs (Tanzania), Santesson (Kenya, Tanzania), Sharp (Tanzania), 
Sipman (Kenya, Tanzania), and Vitikainen (Kenya). Specimens cited with a number but 
without a collector's name were collected by one of the authors. Specimens have been 
deposited in O, BM, and East African herbaria. 

The material referred to in this study has been subjected to thin-layer chromatography 
(TLC) by means of standard methods (C. Culberson, 1972; Menlove, 1974). However, type 
specimens which have recently been tested by TLC have not been retested unless it was 
specially indicated. All specimens cited, including types, have been personally studied 
unless otherwise noted. In the text + denotes a low concentration of a given substance, whilst 
± denotes its sporadic occurrence (accessory substance). 

The descriptions of the species are based on East African material, and comprise only 
those morphotypes and chemotypes which occur in our area. Only selected East African 
records are given for the commoner species, but in such cases at least one record is chosen 
from each district (or, for Uganda, each county) in which the species has been collected. Only 
new or otherwise interesting synonyms are given. For a more complete citation see Hale 
(1965). 



Results 

Morphology 

The apothecia of Parmelia tinctorum, the type species of the subgenus, are substipitate and 
laminal. But in the majority of East African species they were found to be submarginal to 
marginal, with swollen and inflated or more or less constricted but well developed stipes. At 
or near the lobe margin the stipe was sometimes formed by a semitubular fold of a 
convoluted lobe, and in such cases the disc might be eccentrically orientated in relation to its 
stipe. 

In contrast to those of other subgenera in Parmelia, Amphigymnia species often have 
perforated apothecial discs. In some species the perforations were found to have developed at 
an early stage in nearly all the apothecia of a thallus, while in others they were present only 
in some of the apothecia of a given thallus or occurred sporadically and seemingly accident- 
ally in old apothecia. 

The spores were simple and ellipsoid and varied in length from 10 to 40 jum. With some 
individual variation, spore size in the subgenus could be roughly divided into three fairly 
well defined categories: (a) small, thin-walled, 10-20 (22) //m; (b) intermediate, usually 
relatively thick-walled, 20-28 //m; and (c) large, thick-walled, 25^40 //m. In the first group it 
was, to a certain degree, even possible to distinguish between species with spores less than c. 
16 fxm and those 16-20 (22) //m in length. Our own observations and previous reports show 
that the East African species could be divided by spore size into the four groups shown in 
Table 1 . In sorediate species the spores were sometimes poorly developed and on the average 
smaller than in their non-sorediate counterparts. 

The pycnidia were situated in a broad submarginal zone in all primary species, and were 



146 H. KROG & T. D. V. SWINSCOW 

Table 1 Approximate spore sizes in the East African species of Parmelia subgenus 
Amphigymnia. 



10-16 (18) jan 


16-20 (22) urn 


20-28 (30) /an 


25^0 pm 


P. aldabrensis 


P. abessinica 


P. dilatata 


P. amaniensis 


P. cetrata 


P. andina 


P. durumae 


P. bangii 


P. erubescens 


P. austrosinensis 


P. euneta 


P. crinita 


P. eurysaca 


P. cooperi 


P. inexpectata 


P. cristifera 


P. grayana 


P. gardneri 


P. lobulascens 


P. cryptoxantha 


P.jacarandicola 


P. hababiana 


P. nilgherrensis 


P. eciliata 


P. maclayana 


P. hanningtoniana 


P. per lata 


P. subarnoldii 


P. pardii 


P. hololoba 


P. subschimperi 




P. reticulata 


P. leonis 






P. sancti-angelii 


P. mellissii 






P. soyauxii 


P. pigmentifera 






P. stuhlmannii 


P. praesorediosa 






P. subcolorata 


P. pseudocrinita 






P. subisidiosa 


P. rava 






P. sub sump ta 


P. rimulosa 






P. subtinctoria 


P. uberrima 






P. taitae 


P. vivida 






P. tinctorum 


P. zollingeri 






P. ultralucens 








P. xanthina 









also found in this position in a number of secondary species. In modern taxonomic treat- 
ments of parmelioid lichens little emphasis has been placed on the conidia, although their 
shape and size were faithfully recorded by most earlier lichenologists as an important part of 
the descriptions of new species. Within some lichen genera there is obviously little variation 
between the species, but in Parmelia s. lat. the conidia were very variable and are therefore 
treated here in some detail. 

In the Parmeliaceae the conidia are produced endobasidially on conidiophores of the 
bayonet type (Gluck, 1899) (Fig. la). The development of the conidiophores, which has been 
discussed by Vobis (1980), is of little interest in the present study and is not treated here. As 
for the conidia, however, we have found that five common types can be recognized within 
the Parmeliaceae: 

(a) Bifusiform conidia. The term has been applied by, for example, Esslinger (1977) and 
Vobis (1980), to short conidia, usually 4-6 (7) //m long, which are constricted in the middle 
with a swelling near but not at both ends (Fig. lb). Other terms for this type are 
'hantelformig' and 'bisquitformig' (Gluck, 1899) and 'dumb-bell-shaped' (Karnefelt, 1979). 
Within the Parmeliaceae we have observed bifusiform conidia in Cetraria, Cetrelia, 
Hypogymnia, - id Parmelia subgenus Hypotrachyna, but never in subgenus Amphigymnia. 

(b) Sublageniform conidia. The term has been used to denote short, rod-shaped conidia, 
5-8 pm long, with a slight swelling near one end (Fig. lc). Nylander (1885) applied the term 
to the conidia of Parmelia corniculans, and Steiner & Zahlbruckner (in Zahlbruckner, 1926) 
to those of P. procera. The term lageniform means flask-shaped, but since the swelling in this 
type of conidia is subterminal rather than terminal, the term sublageniform is adopted. We 
find this term more appropriate than Gluck's choice 'keulig' or clavate (Gluck, 1899). In 
some species the swelling is indistinct and may be discernible only in a few of the many 
conidia from a pycnidium, while the rest may appear rod-shaped. In the Parmeliaceae sensu 
Henssen & Jahns (1973) sublageniform conidia occur in, for example, Usnea (Swinscow & 
Krog, 1976, Fig. 7), Letharia, Platismatia, and Parmelia subgenus Amphigymnia pro parte. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



147 







Fig. 1 Conidia in genus Parmelia s. lat. a: conidiophore with filiform conidia, P. euneta Stirton, 
3K 16/122. b: bifusiform conidium, P. ducalis Jatta, E 30/12. c: sublageniform conidium, P. 
maclayana Miill. Arg., Ryvarden 9032. d: unciform conidium, P. subpraesignis Nyl., T 16/137. 
e: rod-shaped conidium, P. eciliata (Nyl.) Nyl., 4 May 1978, Dahls.n. 



Sublageniform conidia with one or two darker 'dots' (see below under rod-shaped conidia) 
have been seen in the subgenus Everniiformes. 

(c) Unciform conidia. We have chosen this term to denote rod-shaped conidia c. 5 /im long, 
with a single hook-shaped end (Fig. Id), a type not discussed by Gluck (1899) or Vobis 
(1980). They are particularly characteristic for Parmelia subgenus Parmelia sect. Simplices. 
occurring in all the species examined by us, with the exception of P. microsticta Miill. Arg. 
and P. hypoleucites Nyl 5. str. in which they were filiform (see also W. Culberson & C. 
Culberson, 1980). The presence of unciform conidia in Amphigymnia has not been con- 
clusively proved. 

(d) Rod-shaped (bacilliform) conidia. This is a rather heterogeneous group, consisting of 
rods mainly 8-10(12) jum long and about 10-12 times as long as broad (Fig. le). They have 
no swellings, but in some cases one or two darker 'dots' are situated in either a central, 
subapical, or apical position. We have made no effort to distinguish between the various 
types of 'dotted rods' in this work. They are of little interest in Amphigymnia but occur in 
the subgenus Cyclocheila and subgenus Parmelia sections Parmelia and Bicornuta. Plain 
(i.e. non-'dotted') rods, on the other hand, occur in a few Amphigymnia species, such as 
Parmelia eciliata and P. mesotropa. They form a transition between weakly sublageniform 
and filiform conidia. 

(e) Filiform conidia. Filamentous conidia have been described as filiform by, for example, 
Vainio (1890), Moberg (1977), and Vobis (1980). Their diameter is usually about 1 //m or 
less, so that the conidia are at least 15-20 times as long as broad (Fig. la). Plain rod-shaped 
conidia mainly differ from filiform conidia in being broader in relation to their length and 



148 



H. KROG & T. D. V. SWINSCOW 



usually less than 10 //m long. In the Parmeliaceae longly filiform, curved conidia, about 
30 //m long, are known in Parmeliopsis. In Parmelia s. lat. filiform conidia (usually less than 
20 fim long) occur rarely in Simplices and Cyclocheila but are common in Amphigymnia. 
They are mainly straight, but occasionally a conidium may be bent or curved. 

The shape and size of most forms of conidia were found to show very little variation within 
a single species. Filiform conidia were a notable exception, varying considerably in length 
from specimen to specimen of a species, although they were often of fairly uniform size 
within individual pycnidia. 

Important thalline characters in the subgenus Amphigymnia are the presence or absence 
of vegetative propagules, cilia, and maculae, and the colour of the lower cortex. These 
characters have been discussed in detail by Hale (1965). For further comments, see below 
under 'Circumscription of the species' (p. 1 53). 

A number of more or less well established species pairs were present in our area. Table 2 
shows the primary species of such pairs and their sorediate and isidiate counterparts. 

Table 2 Species counterparts in East African representatives of Parmelia subgenus 
Amphigymnia. 



No soralia or isidia 



Sorediate 



Isidiate 



P. abessinica 
P. aldabrensis 
P. amaniensis 
P. andina 
P. cetrata 
P. eciliata 
P. erubescens 
P. euneta 
P. hololoba 
P.jacarandicola 
P. mac lay ana 
P. nilgherrensis 
P. soyauxii 
P. subcolorata 
P. taitae 
P. uberrima 
P. vivida 
P. zollingeri 



P. hababiana 

P. subarnoldii 

P. austrosinensis 

P. reticulata 

P. perlata (?) 

P. sub sump ta 

P. subschimperi 

P. cooperi 

P. direagens (?) 

P. poo Hi 

P. lobulascens 

P. defecta 

P. permutata 

P. pseudograyana 

P. louisianae (?) 

P. aprica 

P. gardneri 



P. kwalensis 



P. subisidiosa 

P. crinita 

P. s ub tine tor ia 

P. pseudocrinita 



P. stuhlmannii 



Chemistry 

Atranorin and usnic acid are present either separately or jointly in the upper cortex. The 
/?-orcinol depsidone psoromic acid is restricted to the soralia in Parmelia direagens, and to 
the cortex near the apothecia in P. jacarandicola. In the medulla the following substances 
occur: orcinol depsides (lecanoric and gyrophoric acids), orcinol depsidones (alectoronic, 
2-collatolic, and physodic acids, norlobaridone, loxodine, and the lividic acid complex), 
/i-orcinol depsides (generally lacking in the species of our area except that the mainly cortical 
substance atranorin may be responsible for the K+ yellow reaction in the medulla of P. 
praesorediosa), ^-orcinol depsidones (stictic, norstictic, salazinic, galbinic, protocetraric, 
fumarprotocetraric, and echinocarpic acids), fatty acids (caperatic and protolichesterinic 
acids, the reddenda fatty acid complex, and other unknowns), a xanthone (lichexanthone), 
and various pigments. The substances, with the species in which they occur, are presented in 
Tables 3-5. Only those of diagnostic value are listed, while substances which in our 



PARMELIA (AMPH/GYMNIA) IN EAST AFRICA 

Table 3 Chemical properties of species without soralia and isidia. 



49 



lee gyr nib ale liv stc nst sal pso pc fpc fat pgm ind atr usn 



P. abessinica 
P. aldabrensis 
P. amaniensis 
P. andina 

P. cetrata 
P. durumae 
P. eciliata 
P. erubescens 

P. euneta 

P. eurysaca 

P. hanningtoniana 

P. hololoba 

P. inexpectata 
P. jacarandicola 
P. leonis 
P. madayana 

P. nilgherrensis 
P. pigmentifera 
P. soyauxii 
P. subcolorata 

P. taitae 
P. uberrima 
P. vivida 
P. zollingeri 



Only substances of diagnostic importance are included, lee = lecanoric acid, gyr = gyrophoric acid, nlb = 
norlobaridone, alc = alectoronic acid (including or-collatolic acid), liv = the lividic acid complex, stc = stictic acid, 
nst = norstictic acid, sal = salazinic acid, pso = psoromic acid, pc = protocetraric acid, fpc = fumarprotocetraric acid, 
fat = fatty acids, pgm = pigments, ind = undetermined substances, atr = atranorin, usn = usnic acid. Symbols: 
x = constant; + = low concentrations; ± = occasionally present (accessory substance) or present in a chemical strain. 



specimens always occur in conjunction with one of the main compounds, such as loxodine, 
oxyphysodic acid, connorstictic acid, constictic acid, unknown with alectoronic acid, and 
unknowns with stictic acid, are omitted from the tables. 

In P. pigmentifera we found two undetermined main substances in the following Rf 
classes: in TDA 2-3 : 4, in HEF 4-5 : 6, and in TA 2-3 : 5-6. The medulla reacted PD+ 
ochraceous, K+ deep yellow, C— , KC+ red. The reactions with K and PD may have been 
caused either by the above mentioned unknowns or by an unknown medullary pigment, 
while the KC reaction points to the presence of one or two orcinol depsidones. Both 
substances gave a bright bluish white fluorescence under long-wave U V light. 

Among the fatty acids of diagnostic importance is an array of substances which occur 
together, having Rf values intermediate between those of caperatic and protolichesterinic 
acids; they are referred to as the 'reddenda fatty acid complex 1 . The name is derived from 
their presence in, for example, P. reddenda Stirton (Krog & Swinscow, 1977). Unidentified 
fatty acids, usually with rather high Rf values, are accessory in some species. 

In the P. dilatata group (including, for example, P. affluent and P. progenes) atranorin and 
usnic acids are produced together in considerable amounts. The proximity of other 
substances sometimes caused the spots indicating usnic acid to take on a dark sordid grey 
colour on the TLC plates. The presence of usnic acid in this species group has gone partly 



150 H. KROG & T. D. V. SWINSCOW 

Table 4 Chemical properties of species with isidia, sorediate isidia, or dactyls. 

lee gyr nib ale phy stc nst sal pc ech fat pgm lex ind atr usn 

P. crinita x x 

P. cryptoxantha xxx.. x 

P. kwalensis xx x 

P. lophogena .x x. .. x 



P. mellissii 
P. planatilobata 
P. pseudochnita 
P. stuhlmannii 



P. subisidiosa x + 

P. subtinctoria .. x .... x ... . .. + 

P. sulphurata x + 

P. tinctorum x x 

P. tsavoensis x x 

P. ultralucens x.... x. x 

P. xanthina x. .x .x 

Only substances of diagnostic importance are included, lee = lecanoric acid, gyr = gyrophoric acid, nlb = 
norlobaridone, ale = alectoronic acid (including cr-collatolic acid), phy = physodic acid, stc = stictic acid, nst = 
norstictic acid, sal = salazinic acid, pc = protocetraric acid, ech = echinocarpic acid, fat = fatty acids, pgm = pigments 
(including vulpinic acid), lex = lichexanthone, ind = undetermined substances, atr = atranorin, usn = usnic acid. 
Symbols: x = constant; + = low concentrations. 

unnoticed, and the grey spots were thought by Hale (1971c, 1977), for instance, to represent 
an unknown substance. 

Usually the chemical properties of lichens remain unchanged after extended storage in the 
herbarium. However, we noticed that in old specimens with alectoronic and a-collatolic 
acids we often obtained additional spots below those of the main substances on the TLC 
plates, correlated with an ochraceous reaction with C in the medulla. They were especially 
prevalent in the P. nilgherrensis group, of which many old collections exist. We found no 
correlation between morphological characters and the presence or absence of such 
substances, and are inclined to regard them as artefacts. On the other hand, an unknown with 
low Rf values, occurring with alectoronic acid in a chemical strain of P. maclayana, appears 
to be of diagnostic value. It will be referred to below as 'unknown with alectoronic acid'. 

The African chemotype of P. xanthina contained an undetermined C— , KC+ red 
substance which is probably close to gyrophoric acid. It ran slightly higher than gyrophoric 
acid on the TLC plates in TDA and HEF. 

Ecology and distribution 

Tropical East Africa offers a wide range of habitats, from the sea coast to more than 5000 m 
altitude, and from subdeserts and savannas to montane rain forests. Very few lichens are 
found in the large subdesert areas of northern Kenya and Uganda and southern Ethiopia, and 
also in the dry thornbush savannas which lie below 1000 m altitude and have an annual 
rainfall of less than 500 mm. 

While species of Parmelia subgenus Hypotrachyna are absent from the East African sea 
coast and lowlands below c. 800 m altitude (Krog & Swinscow, 1979), the subgenus 
Amphigymnia is represented there by more than a dozen species. They grow partly in 
mangroves at sea level and partly in the coastal forest and low coastal hills up to c. 500 m. All 
are drought tolerant, and several of them are found also in the upland savannas. The coastal 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 

Table 5 Chemical properties of species with soralia. 



151 



lee gyr nib ale stc nst sal pso gib pc fpc ech fat pgm atr usn 



P. aprica 
P. araucariarum 
P. austrosinensis 
P. bangii 

P. cooperi 
P. cristifera 
P. defecta 
P. dilatata 

P. direagens 
P. gardneri 
P. grayana 
P. hababiana 

P. indoafra 
P. lobulascens 
P. louisianae 
P. parahypotropa 

P. pardii 
P. perlata 
P.permutata 
P. pilosa 

P. poolii 
P. praesorediosa 
P. pseudo gray ana 
P. rava 

P. reticulata 
P. rimulosa 
P. sancti-angelii 
P. subarnoldii 

P. subschimperi 
P. subsumpta 
P. umbrosa 



x x 

x 
x 



x x 
x 



Only substances of diagnostic importance are included, lee = lecanoric acid, gyr = gyrophoric acid, nlb = 
norlobaridone, ale = alectoronic acid (including a-collatolic acid), stc = stictic acid, nst = norstictic acid, sal = 
salazinic acid, pso = psoromic acid, glb = galbinic acid, pc = protocetraric acid, fpc = fumarprotocetraric acid, 
ech = echinocarpic acid, fat = fatty acids, pgm = pigments, atr = atranorin, usn = usnic acid. Symbols: x = constant; + = 
low concentrations; ± = occasionally present (accessory substance) or present in a chemical strain. 



Table 6 Coastal species ofParmelia subgenus Amphigymnia in East Africa, arranged by the 
lowest altitude at which they occur, and showing the altitudinal range in metres. 



P. kwalensis 





P. gardneri 


0-2000 


P. pigmentifera 


0-70 


P. uberrima 


0-2000 


P. parahypotropa 


0-300 


P. tinctorum 


0-2700 


P. aldabrensis 


0-1000 


P. reticulata 


0-3000 


P. zollingeri 


0-1000 


P. dilatata 


300-900 


P. durumae 


0-1100 


P. cristifera 


300-1450 


P. hololoba 


0-1800 


P. sulphur ata 


480 



152 H. KROG & T. D. V. SWINSCOW 

Table 7 Photophilous, drought tolerant inland species of Parmelia subgenus Amphigymnia 
in East Africa, arranged by the lowest altitude at which they occur, and showing the altitudinal 
range in metres. See Table 6 for additional species. 



P. Icon is 


700-1400 


P. gray ana 


1160-1500 


P. praesorediosa 


700-1800 


P. rava 


1250-1800 


P. hababiana 


800-2650 


P. pilosa 


1300-2000 


P. andina 


900-2400 


P. subtinctoria 


1300-2300 


P. maclayana 


900-2200 


P. pseudo gray ana 


1400-1900 


P. pool ii 


900-2300 


P. indoafra 


1450-2600 


P. abessinica 


900-2600 


P. xanthina 


1500 


P. vivid a 


920 


P. stuhlmannii 


1500-2100 


P. aprica 


1000 


P. coo peri 


1 500-3000 


P. louisianae 


1000 


P. pardii 


1650-1750 


P. tsavocnsis 


1000 


P. hanningtoniana 


1700 


P. soyauxii 


1000-2100 


P. defecta 


1750-2100 


P. sancti-angelii 


1000-2800 


P. jacarandicola 


1900 


P. austrosinensis 


1000-3000 


P. taitae 


1980-2050 



species, with their altitudinal ranges, are shown in Table 6; they include, for example, the 
widespread, photophilous P. tinctorum and the ubiquitous P. reticulata. 

Drought tolerant photophilous species which do not reach the coast are found in savannas, 
bushed grassland, well lit sites in the montane forests, and artificial habitats such as parks, 
plantations, town avenues, and roadside trees and gardens (Table 7). 

Species with a somewhat higher moisture requirement are found in the more shaded sites 
in the lower montane forests and those parts of the inselbergs which are regularly influenced 
by mist. They often grow together with savanna species, but are here included among those 
of the montane forests when they are largely absent from drier habitats. Species that are more 
exacting in their moisture requirements are often best developed in the upper montane forest 
where the rainfall is higher and more equally distributed throughout the year, especially on 
the wetter slopes of the mountains. In the lower part of the alpine zone, above c. 3500 m, 
there is still ample moisture, combined with high light intensity and low night temperatures. 
Only four of our species, P. euneta, P. lobulascens, P. nilgherrensis, and P. subschimperi, 
extend through the ericaceous zone and into the low alpine zone. 



Table 8 Montane and low alpine species of Parmelia subgenus Amphigymnia in East Africa, 
arranged by the lowest altitude at which they occur, and showing the altitudinal range in 
metres. Parmelia reticulata is an additional species, included in Table 6. 



P. subarnoldii 


500-3000 


P. crinita 


1400-3400 


P. pseudocrinita 


700-2600 


P. direagens 


1500-2600 


P. planatilobata 


800-1000 


P. bangii 


1 500-3200 


P. amaniensis 


800-1150 


P. cryptoxantha 


1550-2100 


P. eciliata 


800-1200 


P. euneta 


1600-3750 


P. ultralucens 


900 


P. subcolorata 


1700-2500 


P. inexpectata 


1100 


P. rimulosa 


1 800-2900 


P. permulaia 


1100-2400 


P. subisidiosa 


1800-3200 


\ umbrosa 


1300-2100 


P. lobulascens 


1800-3600 


P. erubescens 


1400 


P. subschimperi 


1800-3600 


P. sub sump la 


1400-2100 


P. araucariarum 


2000-2100 


P. lophogena 


1400-2400 


P. mellissii 


2000-2400 


P. celrata 


1400-2600 


P. nilgherrensis 


2000-3600 


P. per lata 


1400-3100 







PARMEL1A (AMPHIGYMNIA) IN EAST AFRICA 

Table 9 Species of Parmelia subgenus Amphigymnia which are not known outside 
Africa. 



153 



P. aldabrensis 


P. kwalensis 


P. stuhlmannii 


P. amaniensis 


P. leonis 


P. subcolorata 


P. aprica 


P. lophogena 


P. subschimperi 


P. cryptoxantha 


P. pardii 


P. taitae 


P. dejecta 


P. pigmentifera 


P. tsavoensis 


P. durumae 


P. pseudograyana 


P. uberrima 


P. hololoba 


P. rava 


P. umbrosa 


P. inexpectata 


P. rimulosa 


P. vivida 


P.jacarandicola 


P. soyauxii 





The species of the montane forests and the lower part of the alpine zone are shown in 
Table 8. (In addition comes P. reticulata, included in Table 6 as one of the species that reach 
the coast.) The subgenus Amphigymnia is not represented in the upper parts of the alpine 
zone, above c. 4000 m altitude. 

Of the 70 Amphigymnia species here recorded for East Africa, 26 species, or nearly 40 per 
cent, are not yet known outside Africa (Table 9). Among the remaining species, 35 are found 
also in the Americas; some of them are so far known only from Africa and the Americas, 
namely Parmelia abessinica, P. araucariarum, P. bangii, IP. eurysaca, P. hanningtoniana, 
IP. louisianae, P. pilosa, and P. subsumpta. Thirty-three are known from Asia, and of these 
species the following are known only from Africa and Asia: Parmelia cooperi, P. direagens, 
IP. euneta, P. indoqfra, P. lobulascens, P. nilgherrensis, P. planatilobata, and P. 
pseudocrinita. Twenty-three of the East African species occur in the Australian region; they 
are all known also from other regions. Only five cosmopolitan, mainly temperate species are 
common to East Africa and Europe; they are P. austrosinensis, P. cetrata, P. crinita, P. 
perlata, and P. reticulata. 



Discussion 
Circumscription of the species 

In his world monograph Hale (1965) discussed the relative taxonomic value of various 
morphological and chemical characters on the basis of their degree of association with each 
other. He found that cilia had a positive or negative correlation with more than half of the 
characters considered, and deduced that they were of importance at species level. Similarly, 
maculae, a white marginal zone on the under side, an entirely brown under side, and a 
perforated apothecial disc were found to be valid characters, whereas spore size and 
ornamentation of the thalline exciple were not. Conidia were assumed to have no 
importance and were not investigated. Chemical strains were accepted in a limited way; 
many of them have been regarded as species in subsequent publications by the same author. 
It should be borne in mind that Hale's monograph appeared before the advent of TLC, so 
that knowledge of the chemical properties of the species was then in many cases inferior to 
that reached during the last decade. 

We have had the advantage of studying the East African Amphigymnias in the field and of 
collecting numerous specimens of the commoner species from sites with differing ecology. 
This has enabled us to evaluate the morphological variation within the species in the light of 
environmental factors, and to form an opinion on the relative taxonomic value of the various 
characters as expressed in the species of our area. As might be expected, we have observed 
that species with a great ecological amplitude usually show a much greater morphological 
variation than species with a limited range. In a moist, shady habitat in an area with favour- 
able temperatures the thallus is often membranaceous, greenish in colour, has long, slender 



154 H. KROG & T. D. V. SWINSCOW 

cilia and distinct maculae, whereas plants growing in a well lit site, subjected to drought or 
low temperatures conducive to slow growth, in general have a coriaceous thallus, a grey 
colour more or less tinged with brown, short, stunted and coarse cilia, and less conspicuous 
maculae. At their extremes such plants look very different and have often been interpreted as 
distinct species. 

Taking the environmentally induced variation into account, we find that most of the 
characters listed by Hale (1965) are of value at species level, but with certain reservations. A 
normally ciliate species, such as Parmelia nilgherrensis, may rarely lack cilia, while a few 
normally eciliate species, such as P. cristifera and P. gardneri, may produce rudimentary 
cilia in the lobe axils. A strongly maculate species clearly differs from a species in which 
maculae are always indistinct or absent, but in a normally maculate species it is possible for 
saxicolous specimens with a coriaceous thallus to develop very few maculate areas. Presence 
or absence of a perforated apothecial disc is fairly constant in most species but varies greatly 
in some. In P. euneta and P. maclayana, for example, the apothecia may all be perforate, or 
all imperforate, or both types may be present on a single specimen, so that the character is of 
limited diagnostic value in these two species. A white marginal zone on the under side is 
characteristic of some species, such as P. leonis and P. uberrima, but occurs only in some 
specimens of, for example, P. abessinica and P. andina. In the P. erubescens species 
complex, as here delimited, the colour of the under side varies from black through brown to 
ivory. 

The various types of vegetative propagules are usually considered to be species specific, 
but in some species there is a complete intergradation between granular soredia, sorediate 
isidia, non-sorediate isidia, and dactyls. This is the case in, for example, P. lophogena and P. 
mellissii. We have also found that P. tinctorum is subject to variation in its vegetative 
propagules, which range from simple or branched, slender, cylindrical isidia to coarse, 
branched isidia and closed or open dactyls. Both isidia and dactyls may at times disintegrate 
partly or wholly into granular soredia. We have chosen to regard P. pseudotinctorum (coarse 
isidia and dactyls) as conspecific with P. tinctorum (slender isidia), since the variation is 
largely continuous and both morphotypes have the same type of spores and conidia, and 
mainly the same ecological requirements. If a distinction were to be maintained, we believe 
that it should be at infraspecific level. 

We do not agree with Hale (1965) that spore size alone is not a valid character. In our 
experience there is little variation in spore length within species whose spores are below 
20 jum long, so that two subgroups whose spores are respectively above and below about 
16 [im can be recognized (Table 1). In species with medium sized spores, on the other hand, 
the spores may vary between 20 and 30 //m, often with a maximum around 22-25 //m, while 
in those with large spores, the spores may in extreme cases show variation between 25 and 
40 jum, usually with a maximum around 28-30 /im. There is in our area a clear discontinuity 
between species with small spores and those with large spores, and we regard this character as 
species specific. 

There is at present no common agreement on the function of the lichen conidia. A sexual 
function for them has not been conclusively proved, but nor has their asexual nature been 
firmly established. Regardless of their function, their constancy within morphologically well 
delimited species and partly also within supraspecific units show that a taxonomic value 
should be assigned to them that would normally not be bestowed upon vegetative 
propagules. In agreement with, for example, Henssen & Jahns (1973) we consider the shape 
of the conidia to be species specific, and like Moberg (1977) we believe that they are of 
taxonomic significance also at the supraspecific level. 

In Amphigymnia we have noted a strong correlation between imperforate apothecia and 
sublageniform conidia on the one hand and perforate apothecia and filiform conidia on the 
other, although there are exceptions to this rule (Table 10). Parmelia abessinica, for 
example, has perforate apothecia but weakly sublageniform conidia. In P. maclayana, in 
which the apothecia may be perforate or not, the conidia are sublageniform, whereas in P. 
euneta, with similarly variable apothecia, they are filiform. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 

Table 10 Number of species with either filiform or 
sublageniform conidia, arranged by presence or absence 
of perforate apothecial discs. 

Conidia 



155 



Apothecia 



filiform 



sublageniform 



perforate 
imperforate 



24 



4 
13 



Table 11 Epithets applied to various chemotypes in the Parmelia 
euneta — Parmelia subschimperi species pair. 



Substances 


No soralia or isidia 


Soralia 


gyrophoric acid 


euneta 

(1877-78) 


neolobulascens 
(1975) 


gyrophoric acid 
norstictic acid 


compos it a 
(1972) 


subcompositum 
(1977) 


gyrophoric acid 
norlobaridone 
norstictic acid (±) 


unnamed 


balensis 
(1975) 


norlobaridone 


unnamed 


subschimperi 
(1972) 


norlobaridone 
norstictic acid 


spilota 
(1973) 


unnamed 


norstictic acid 


unnamed 


— 



In the delimitation of the species we assign greater taxonomic value to characters of spores 
and conidia than we do to characters of the thallus. Many secondary species without 
apothecia still produce pycnidia, and if conidia can be found they offer a valuable guide to 
the correct interpretation of the species. 

Chemical characters show no continuous variation, with the exception of accessory 
substances which vary from abundance to absence within a given species and are commonly 
regarded as of no taxonomic value. All other cases of chemical variation represent easily 
registered discontinuities, and have therefore been regarded as species specific by some 
authors. However, as with the morphological variation we have attempted to evaluate each 
case of chemical variation individually on the basis of trends observed in our material. 

Winnem (1975) clarified the chemical properties of the sorediate species P. direagens. She 
found that it had three chemical strains: (1) psoromic acid, gyrophoric acid, and alectoronic 
acid; (2) psoromic acid and gyrophoric acid; and (3) psoromic acid and alectoronic acid. The 
psoromic acid, an extremely rare substance in Amphigymnia, is mainly connected with the 
soralia in P. direagens, which is an unusual situation in Parmelia but known in certain other 
genera, for example Usnea (Swinscow & Krog, 1979). In our opinion the presence of this 
common denominator unites the morphologically uniform chemotypes and shows beyond 
doubt that they merely represent chemical strains within a species. Apart from the presence 
of psoromic acid, a depside and a depsidone of the orcinol series occur separately or jointly. 

A parallel to this example is found in the chemical variation of the P. euneta — P. 
subschimperi species pair, although there is no common denominator present (Table 1 1). 



156 



H. KROG & T. D. V. SWINSCOW 



The species comprise strongly maculate, ciliate plants with intermediate spores, filiform 
conidia, and a montane to low alpine distribution. Winnem (1975) showed that norstictic 
acid was accessory in this group and of no taxonomic importance. The remaining substances, 
gyrophoric acid and norlobaridone, may occur separately or jointly. As in P. direagens this is 
also a situation involving a depside and a depsidone of the orcinol series. 

The P. amaniensis — P. subarnoldii pair are ciliate, emaculate species with imperforate 
apothecia, a strongly dentate-ciliate thalline margin, large, thick-walled spores, and 
sublageniform conidia. The following chemical strains occur in each species: (1) 
protocetraric acid and alectoronic acid; (2) protocetraric acid and a-collatolic acid, ± 
alectoronic acid; and (3) protocetraric acid and protolichesterinic acid (Table 12). Spores, 
conidia, and thallus morphology afford several valid characters by which to define this 
species pair, and chemically protocetraric acid is a common denominator. We have here a 
substitution situation involving two closely related orcinol depsidones and a fatty acid. 

Another example involving an orcinol depsidone and a fatty acid is that of the P. 
abessinica — P. hababiana species pair, each of which is known in the following chemical 
strains: (1) norlobaridone; (2) norlobaridone and protolichesterinic acid; and (3) proto- 
lichesterinic acid. 

A group of mainly Australian and South American species poses problems related to 
variation in both cortical and medullary substances. All specimens have imperforate or 
narrowly perforate apothecia, small spores, and a distinctly maculate upper side. The under- 
side varies from black in the centre with a broad, brown or mottled marginal zone through 
uniformly brown to ivory. The rhizines are usually dimorphous, and rhizines and papillae 
often, but not always, reach the lobe margins. The majority of species have negligible 
quantities of atranorin in the cortex, often in too small amounts to be detected with TLC, but 



Table 1 2 Epithets applied to various chemotypes in the Parmelia 
amaniensis — Parmelia subarnoldii species pair. 



Substances 


No soralia or isidia 


Soralia 


protocetraric acid 
alectoronic acid 


amaniensis 
(1926) 


deflectens 
(1979) 


protocetraric acid 
(2-collatolicacid 


pach vspora 
(1965) 


unnamed 


protocetraric acid 
protolichesterinic acid 


unnamed 


subarnoldii 
(1961) 



Table 13 The oldest epithets for the various chemotypes in the Parmelia 
erubescens species complex. 



Substances 


No soralia or 


isidia 


Sorediate 


Isidiate 


salazinicacid 


erubescens 
(1877-78) 




subsumpta 
(1869) 


unnamed 


norlobaridone 


recipienda 




confer endum 


haitiensis 




(1885) 




(1977) 


(1959) 


salazinicacid 


unnamed 




reitzii 


subtinctoria 


norlobaridone 






(1977) 


(1930) 



PARMEL/A (AMPHIGYMNIA) IN EAST AFRICA 157 

some Central and South American species, such as P. subcaperata Krempelh. and 
Parmotrema neotropicum Kurok., have copious amounts of usnic acid together with 
medium amounts of atranorin. In the Parmeliaceae there is no precedent for accepting such 
large variation in cortical substances within a species. The presence of usnic acid in the 
cortex is here correlated with salazinic acid in the medulla, while species with only traces of 
atranorin in the cortex may produce salazinic acid and norlobaridone separately or jointly. 
That the variation in cortical substances does not parallel that of the medulla is a further 
reason for regarding the species that produce usnic acid as distinct from those with only 
traces of atranorin (referred to below as the P. erubescens complex). 

The oldest epithets available for each chemotype within the P. erubescens species 
complex are listed in Table 13, providing an example of chemical variation involving an 
orcinol depsidone and a /?-orcinol depsidone. As in the examples mentioned above we 
believe that these chemotypes also should be regarded as chemical strains. However, since 
the relevant primary species are poorly represented in our area, they are in need of further 
study, especially with regard to their conidial characters, in order to ascertain their relation- 
ship with each other. We have therefore chosen to treat the chemotypes occurring in our area 
as chemical strains, but do not formally reduce the extra-African chemotypes to synonymy 
at this point. 

When the chemical relationship seemed more complicated than in the examples given 
above, or we felt that we had studied insufficient material, we have hesitated to apply the 
theory of chemical strains until further evidence can be produced. Some examples follow. 

The morphological differences between the two species pairs P. euneta — P. subschimperi 
and P. nilgherrensis — P. lobulascens are few and insignificant; they are mainly that the P. 
nilgherrensis pair has a higher frequency of perforate apothecia and on the whole a more 
coriaceous thallus and more irregularly incised lobes. However, the two species pairs differ 
both in basic chemistry (gyrophoric acid and norlobaridone versus alectoronic acid) and in 
accessory substances (norstictic acid versus gyrophoric acid), while we have not yet come 
upon specimens with a combination of the two orcinol depsidones alectoronic acid and 
norlobaridone. In view of these considerable chemical differences, which are correlated with 
a few weak morphological traits, we regard the two species pairs as distinct from each other. 

Parmelia inexpectata differs from P. amaniensis in that it lacks protocetraric acid (the 
common denominator in the P. amaniensis complex), while it produces an array of orcinol 
depsidones (lividic acid and associated substances), none of which occur in P. amaniensis. 
Since P. inexpectata is known from very few collections, we feel that we have insufficient 
grounds for including the species in P. amaniensis for the time being, especially since it 
differs in several chemical characters. 

The two sorediate species P. grayana (protolichesterinic acid) and P. pseudograyana 
(fumarprotocetraric and protocetraric acids) are morphologically virtually indistinguishable. 
The supposed parent morph of P. pseudograyana, P. taitae, is known only with the ^-orcinol 
depsidones. This species pair appears to be restricted to Africa, while P. grayana is widely 
distributed. We should like to see much more material of P. taitae, and preferably also fruit 
bodies in P. pseudograyana, before we make a final decision on the relationship between the 
two secondary species. 

It is difficult to assess the relationship between chemically discordant secondary species 
when the corresponding primary species are not known. In such cases it is preferable to 
regard the species as distinct until further evidence becomes available. 

The underlying causes of chemical diversity in lichens are not known. Theoretically, a 
single mutation influencing the biogenetic pathway at an early point may change the 
chemical end product considerably. If sexual reproduction is assumed to take place among 
lichens, cross fertilization between two individuals with different chemical properties might 
result in the spores from one apothecium giving rise to plants of different chemotypes. If one 
further assumes that some gene exchange takes place, a new chemotype characterized by a 
combination of substances may result. Chemotypes originating in that manner would 
probably not be regarded as distinct species. 



158 H. KROG & T. D. V. SWINSCOW 

It is possible that conidia rarely function as spermatia in lichens, or, if they do, that 
fertilization usually takes place between conidia and ascogones produced on the same 
thallus. However, if cross fertilization does occur under certain circumstances, the 
conditions in the tropical rain-forests would be conducive to it. There the various lichens are 
crowded and often multilayered on tree trunks and branches, and they are subjected almost 
daily to tropical showers and water condensed from heavy mist, which could carry conidia 
from one plant to another with minimal risk of desiccation. 



Generic concept and infrageneric classification 

Genus Parmelia s. lat. has in recent years been divided into a number of genera. As we have 
stated earlier (Krog & Swinscow, 1979), we believe some of these genera to be based on 
characters of little importance at the generic level, the result being obscure delimitation 
between some neighbouring genera. We have therefore been reluctant to accept the new 
genera proposed, although we believe that a satisfactory division of Parmelia s. lat. into 
several genera will eventually be reached. 

According to Hale's main division of Parmelia s. lat., Parmelia s. str. has a paraplec- 
tenchymatous upper cortex and a non-pored epicortex frequently bearing pseudocyphellae, 
while the other segregates possess a palisade plectenchymatous upper cortex and a pored 
epicortex without pseudocyphellae (Hale, 1976a). We agree in this major division, but have 
not yet reached a decision on the interpretation of the residual genera in Parmelia s. lat. 
Although the generic name Parmotrema primarily applies to the group of lichens that are 
dealt with here, we hesitate to adopt it until the genus has been satisfactorily circumscribed, 
notably in relation to Parmelia subgenus Cyclocheila sensu Krog & Swinscow (1979). The 
conidia may prove to be of considerable value in the circumscription of the parmelioid 
genera. 

For a subdivision of Amphigymnia, various characters have been employed. Vainio (1890) 
relied on the colour of the upper cortex (presence or absence of usnic acid) when he proposed 
the subsections Subflavescens Vainio and Subglaucescens Vainio. Gyelnik (1932) regarded 
Vainio's subsections as sections and further subdivided them into each two subsections 
(Eciliatae — Ciliatae, Eciliolae — Ornaticolae) on the basis of presence or absence of cilia. 

Hale (1965) found that the presence or absence of cilia was a character of greater 
importance at section level than the colour of the upper cortex. He proposed the following 
scheme: 

Sect. Amphigymnia [lobe margins eciliate or rarely with sparse cilia in the axils]. 
Sect. Subflavescentes (Vainio) Gyelnik [margins and apices of lobes distinctly ciliate]. 
Subsect. Subflavescentes [upper cortex emaculate]. 
Ser. Subflavescentes [usnic acid present]. 
Ser. Emaculatae Hale [usnic acid absent]. 
Subsect. Ornaticolae Gyelnik [upper cortex maculate]. 
Ser. Subpallidae Hale [under side brown, rhizinate to the margins]. 
Ser. Ornaticolae [under side black in the centre, with a commonly white or mottled, 
naked marginal zone]. 

We are undecided whether to regard the presence of cilia as a valid character for infrageneric 
classification in Amphigymnia, since its correlation with other characters is not yet clear. In 
ser. Subpallidae there appears to be correlation between a strongly maculate upper cortex, 
rhizine morphology, and chemical traits, whilst in most other cases the presence of maculae 
seems uncorrelated with other characters. We find the presence or absence of usnic acid in 
the cortex to be of no taxonomic value above species level. However, a classification of the 
amphigymnioid species can only be settled on the basis of a world wide study. Before that is 
possible, the delimitation of the genus Parmotrema must be conclusively agreed upon. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 159 

The East African Amphigymnia species 



Key to the species 



la Soralia and isidia absent 2 (p. 1 59) 

lb Soralia or isidia present 46 

46a Isidia (including sorediate isidia and dactyls) present , 47 (p. 161) 

46b Isidia absent. Soralia present 62 (p. 162) 

Soralia and isidia absent 

2a (la) Marginal cilia absent 3 

2b Marginal cilia present 6 

3a (2a) Thallus yellow (usnic acid present) 68. P. vivida (p. 223) 

3b Thallus grey (usnic acid absent) , 4 

4a (3b) Apothecia imperforate. Medulla C— , PD+ orange-red (protocetraric acid) 

70. P. zollingeri (p. 225) 
4b Apothecia perforate. Medulla C+ red, PD— (lecanoric acid) 5 

5a (4b) Plant saxicolous, strongly attached. Lobes less than 1 cm broad 53. P. soyauxii (p. 2 1 0) 
5b Plant normally corticolous, loosely attached. Lobes more than 1 cm broad 4. P. andina (p. 168) 

6a (2b) Medulla pigmented pale yellow, ochraceous, or salmon pink, pigment K— ... 7 
6b Medulla white, at most with patches of an ochraceous, K+ purple pigment near the 

lower cortex 8 

7a (6a) Apothecia imperforate, with a dentate-ciliate thalline margin. Medulla UV+ 

(undetermined substances). Coastal species .... 42. P. pigmentifera (p. 201) 

7b Apothecia perforate, with a smooth, eciliate thalline margin. Medulla UV— 

(gyrophoric acid). Upland species 56. P. subcolorata (p. 213) 

8a (6b) Upper cortex with a reticulate pattern of maculae and cracks. Rhizines in part 

squarrose. Salazinic acid present 9. P. cetrata(p. 172) 

8b Upper cortex without a reticulate pattern of maculae and cracks. Rhizines not 

squarrose. Salazinic acid present or absent • 9 

9a (8b) Underside with a distinct, white marginal zone 10 

9b Underside with a brown or mottled marginal zone 14 

10a (9a) Apothecia perforate. Medulla UV— .11 

10b Apothecia mainly imperforate. Medulla UV+ 13 

11a (10a) Underside almost entirely white. Medulla C— , KC— (protolichesterinic acid) 

31. P. leonis(p. 191) 
lib Underside black in the centre, white peripherally. Medulla C+ or C— , KC+ or KC— . 12 

1 2a (lib) Lobe margins flat or revolute. Medulla C+ red (lecanoric acid) 26. P. hololoba (p. 1 86) 
12b Lobe margins ascending. Medulla C— (norlobaridone and/or protolichesterinic acid) 

1. P. abessinica (p. 165) 

13a (1 0b) Spores less than 20 /urn long. Only alectoronic acid present 65. P. uberrima (p. 220) 

1 3b Spores more than 20 /xm long. Both alectoronic and or-collatolic acids present 

17. P. durumae (p. 178) 

14a (9b) Apothecia present 15 

14b Apothecia absent .30 



160 H. KROG & T. D. V. SWINSCOW 

15a ( 14a) Spores more than 20 /im long 16 

15b Spores less than 20 jum long 21 

16a ( 15a) Upper cortex emaculate. Apothecia imperforate 17 

16b Upper cortex distinctly maculate. Apothecia perforate or imperforate 20 

17a (1 6a) Apothecia with an even, eciliate thalline margin. Stictic acid present 

18. P. eciliata (p. 180) 
1 7b Apothecia normally with a dentate-lobulate and/or ciliate thalline margin. Stictic acid 

absent -18 

1 8a (1 7b) Spores 22-26 //m long. Alectoronic and or-collatolic acids present 17. P. durumae (p. 1 78) 
18b Spores 25-28 (40) //m long. Chemistry various • 19 

19a (18b) Medulla with substances in the lividic acid complex . . . 28. P inexpectata (p. 188) 
19b Medulla with protocetraric acid (usually in combination with alectoronic, <z-collatolic, 

or protolichesterinic acid) 3. P. amaniensis (p. 167) 

20a ( 1 6b) Medulla U V+ (alectoronic acid, ± <2-collatolic acid, ± gyrophoric acid) 

37. P. nilgherrensis (p. 197) 
20b Medulla UV— (gyrophoric acid and/or norlobaridone, ± norstictic acid) 20. P. euneta (p. 1 8 1 ) 

21a (15b) Upper cortex distinctly maculate 22 

21b Upper cortex emaculate or faintly maculate 25 

22a (21a) Thalline exciple with coarse, isidioid protuberances. Medulla C+ rose, PD— 

(gyrophoric acid) 25. P. hanningtoniana (p. 185) 

22b Thalline exciple smooth to rugose, without isidioid protuberances. Medulla C— , PD+ 

orPD- 23 

23a (22b) Rhizines dimorphous. Salazinic acid and norlobaridone present 

19. P. erubescens (p. 180) 
23b Rhizines uniform. Chemical properties otherwise 24 

24a (23b) Underside brown. Medulla PD+ orange (stictic and norstictic acids). Coast and 

lowland species 2. P. aldabrensis (p. 166) 

24b Underside black in the centre, white, mottled, or brown peripherally. Medulla PD— 

(norlobaridone and/or protolichesterinic acid). Upland species 1 . P. abessinica (p. 1 65) 

25a (21b) Cortex PD+ sulphur yellow near the apothecia (psoromic acid present). Medulla 

C+ rose (gyrophoric acid) 29. P. jacarandicola (p. 190) 

25b Cortex at most PD+ pale yellow (psoromic acid absent). Medulla C+ or C— . ... 26 

26a (25b) Medulla UV+ (alectoronic acid) 35. P. maclayana (p. 195) 

26b Medulla UV— (alectoronic acid absent) 27 

27a (26b) Plant saxicolous. Apothecia imperforate. Medulla PD+ orange-red (fumarproto- 

cetraric acid) 62. P. taitae(p. 217) 

27b Plant normally corticolous. Apothecia perforate. Medulla PD+ or PD— (fumarproto- 

cetraric acid absent) 28 

28a (27b) Central lobes laciniate. Medulla PD+ orange (salazinic acid) 21 . P. eurysaca (p. 1 82) 
28b Central lobes not laciniate. Medulla PD- 29 

29a (28b) Lobe margins flat or revolute. Medulla C+ red (lecanoric acid) 26. P. hololoba (p. 1 86) 
29b Lobe margins ascending. Medulla C— (norlobaridone and/or protolichesterinic acid) 

(norlobaridone and/or protolichesterinic acid). Upland species 1 . P. abessinica (p. 1 65) 

30a ( 14b) Medulla PD+ orange or red . 31 



P ARM ELI A (AMPHIGYMNIA) IN EAST AFRICA 161 

30b Medulla PD— (but cortex may be PD+ sulphur yellow, see 42b) 37 

31a (30a) Plant saxicolous. Fumarprotocetraric acid present . . 62. P. taitae (p. 2 1 7) 
31b Plant normally corticolous. Fumarprotocetraric acid absent 32 

32a (3 lb) Upper cortex distinctly maculate 33 

32b Upper cortex emaculate or faintly maculate . 35 

33a (32a) Salazinic acid and norlobaridone present ... 19. P. erubescens (p. 1 80) 
33b Salazinic acid absent, norlobaridone present or absent . 34 

34a (33b) Montane forest species. Norstictic acid present in combination with gyrophoric 

acid and/or norlobaridone 20. P. euneta(p. 181) 

34b Coastal species. Norstictic acid present in combination with stictic acid 

2. P. aldabrensis (p. 166) 

35a (32b) Protocetraric acid present (usually in combination with alectoronic, or-collatolic, 

or protolichesterinic acid) 3. P. amaniensis (p. 167) 

35b Protocetraric acid absent 36 

36a (35b) Salazinic acid present 21. P. eurysaca(p. 182) 

36b Stictic acid present 18. P. eciliata(p. 180) 

37a (30b) Upper cortex distinctly maculate • 38 

37b Upper cortex emaculate or faintly maculate 41 

38a (37a) Species of dry, well lit upland habitats . 39 

38b Species of the montane forests and the alpine zone .40 

39a (38a) Medulla C+ rose (gyrophoric acid) .... 25. P. hanningtoniana (p. 185) 

39b Medulla C— (norlobaridone and/or protolichesterinic acid) 1. P. abessinica (p. 165) 

40a (38b) Medulla UV+ (alectoronic acid, ± <2-collatolic acid, ± gyrophoric acid) 

37. P. nilgherrensis (p. 197) 
40b Medulla UV— (gyrophoric acid and/or norlobaridone) ... 20. P. euneta (p. 181) 

41a (37b) Medulla C+ rose or red * 42 

41b Medulla C- . . , 43 

42a (41 a) Lecanoric acid present 26. P. hololoba(p. 186) 

42b Gyrophoric acid present (combined with psoromic acid in the cortex of some lobes) 

29. P.jacarandicola(p. 190) 

43a (4 1 b) Substances in the lividic acid complex present . . 28. P. inexpectata (p. 1 88) 
43b Substances in the lividic acid complex absent 44 

44a (43b) Medulla U V— (norlobaridone and/or protolichesterinic acid) 1 . P. abessinica (p. 165) 
44b Medulla UV+ (alectoronic acid, ± (2-collatolic acid) 45 

45a (44b) Plant corticolous or saxicolous. Upland species above c. 1000 m altitude 

35. P. maclayana (p. 195) 
45b Plant corticolous. Coastal and lowland species below c. 1 000 m altitude 

17.P.durumae(p. 178) 

46a (lb) Isidia (including sorediate isidia) or dactyls present • 47 

46b Isidia and dactyls absent. Soralia present 62 

Isidia (including sorediate isidia and dactyls) present 

47a (46a) Marginal cilia absent . 48 



162 H. KROG & T. D. V. SWINSCOW 

47b Marginal cilia present 51 

48a (47a) Medulla PD+ orange-red (protocetraric acid) 71. P. sp. A (p. 226) 

48b Medulla PD- 49 

49a (48b) Plant large, loosely fastened to tree bark or rock. Cylindrical isidia, granular 

isidia, or dactyls present. Medulla C+ red (lecanoric acid) 63. P. tinctorum (p. 2 1 8) 

49b Plant small, strongly fastened to rock. Only dactyls or coarse isidia present. Medulla C+ 

orC- 50 

50a (49b) Medulla C+ red (lecanoric acid) 54. P. stuhlmannii (p. 21 1) 

50b Medulla C- (physodic acid) 64. P. tsavoensis (p. 220) 

51a (47b) Thallus yellow (usnic acid present) 69. P. xanthina (p. 224) 

51b Thallus grey (usnic acid absent) 52 

52a (51b) Upper cortex with a reticulate pattern of maculae and cracks. Rhizines in part 

squarrose 57. P. subisidiosa (p. 214) 

52b Upper cortex without a reticulate pattern of maculae and cracks. Rhizines not squarrose 53 

53a (52b) Medulla pigmented pale to bright yellow or orange 54 

53b Medulla white, at most with patches of an ochraceous, K+ purple pigment near the 

lower cortex 55 

54a (53a) Medulla pigmented bright yellow to orange throughout. Cylindrical isidia present. 

Coastal species 61. P. sulphurata (p. 217) 

54b Medulla pigmented pale yellow, pigment sometimes inapparent. Open dactyls present. 

Lower montane forest species 13. P. cryptoxantha (p. 175) 

55a (53b) Isidia flattened, dorsiventral. Gyrophoric acid present 44. P. planatilobata (p. 203) 
55b Isidia cylindrical. Chemistry various 56 

56a (55b) Upper side distinctly maculate 57 

56b Upper side emaculate or faintly maculate 58 

57a (56a) Rhizines dimorphous. Salazinic acid and norlobaridone present. Upland species 

60. P. subtinctoria (p. 2 1 6) 
57b Rhizines uniform. Stictic and norstictic acids present. Coastal species 30. P. kwalensis (p. 191) 

58a (56b) Thallus coriaceous. Isidia mainly laminal, never becoming sorediate . . . 59 

58b Thallus usually membranaceous. Isidia mainly submarginal, often becoming sorediate- 

granular 60 

59a (58a) Medulla C+ rose, PD- , UV- (gyrophoric acid) . 47. P. pseudocrinita (p. 205) 

59b Medulla C— , PD+ orange, UV+ intensely yellow (salazinic acid, lichexanthone) 

66. P. ultralucens (p. 221) 

60a (58b) Upper cortex usually continuous. Isidia rarely sorediate. Medulla PD+ orange 

(stictic acid) v 11. P. crinita(p. 173) 

60b Upper cortex fragile and flaking. Isidia often sorediate. Medulla PD— 61 

61a (60b) Medulla C+ rose, UV— (gyrophoric acid) 33. P. Iophogena(p. 194) 

61b Medulla C-,UV+(alectoronic acid) 36. P. mellissii (p. 197) 

Isidia absent. Soralia present 

62a (46b) Marginal cilia absent 63 

62b Marginal cilia present 73 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 163 

63a (62a) Plant saxicolous 64 

63b Plant corticolous 67 

64a (63a) Medulla PD+ orange-red (protocetraric acid) 39. P. pardii (p. 199) 

64b Medulla PD- 65 

65a (64b) Medulla C- (fatty acids) 46. P. praesorediosa (p. 205) 

65b Medulla C+ red (lecanoric acid) '. 66 

66a (65b) Thallus small, coriaceous, strongly attached. Upper cortex emaculate, shiny 

14. P. defecta (p. 176) 
66b Thallus usually large, relatively thin, loosely attached. Upper cortex faintly maculate, 

often matt 7. P. austrosinensis (p. 171) 

67a (63b) Thallus yellow or yellowish grey (usnic acid present). Medulla PD+ orange-red 

(protocetraric acid) 68 

67b Thallus pale grey (usnic acid absent). Medulla PD+ or PD— 70 

68a (67a) Thallus bright yellow to yellow-green. Atranorin absent . . . 5. P. aprica (p. 1 69) 
68b Thallus yellowish grey. Atranorin present . 69 

69a (68b) Echinocarpic acid and various unknowns present ... 15. P. dilatata (p. 1 77) 
69b Echinocarpic acid and unknowns absent 49. P. rava (p. 207) 

70a (67b) Medulla C+ red (lecanoric acid) .... 7. P. austrosinensis (p. 171) 
70b Medulla C- 71 

71a (70b) Medulla PD- (fatty acids) 46. P. praesorediosa (p. 205) 

71b Medulla PD+ orange or red 72 

72a (7 lb) Medulla K+ red (salazinic acid) 12. P. cristifera (p. 173) 

72b Medulla K+ sordid brown (protocetraric acid) ..... 22. P. gardneri(p. 183) 

73a (62b) Upper cortex fragile and flaking. Soralia erupting in a pustular fashion or more or 

less mixed with isidia 74 

73b Upper cortex continuous. Soralia rarely pustular, isidia absent 77 

74a (73a) Medulla pigmented pale yellow. Echinocarpic acid and fatty acids present 

13. P. cryptoxantha (p. 1 75) 
74b Medulla white. Chemistry otherwise 75 

75a (74b) Medulla PD+ orange, UV- (stictic acid) ...... 8. P. bangii (p. 171) 

75b Medulla PD— , UV+(alectoronic acid) 76 

76a (75b) Soralia pustular, without isidia 51. P. rimulosa(p. 209) 

76b Soralia not pustular, but here and there interspersed with isidia . 36. P. mellissii (p. 197) 

77a (73b) Medulla pigmented yellow, salmon pink, or ochraceous, pigment K— . ... 78 
77b Medulla white, at most with patches of an ochraceous, K+ purple pigment near the 

lower cortex 79 

78a (77a) Medulla C+ red in upper parts (gyrophoric acid) .... 41. P. permutata (p. 201) 
78b Medulla C— (fatty acids) 6. P. araucariarum (p. 170) 

79a (77b) Upper cortex with a reticulate pattern of maculae and cracks. Rhizines in part 

squarrose 50. P. reticulata (p. 208) 

79b Upper cortex without a reticulate pattern of maculae and cracks. Rhizines not squarrose 80 

80a (79b) Underside with a distinct, white marginal zone 81 



164 H. KROG & T. D. V. SWINSCOW 

80b Underside with a brown or mottled marginal zone 84 

81a (80a) Lobes deeply divided, with sublinear laciniae. Medulla K+ red, PD+ orange 

(norstictic, galbinic, and salazinic acids). Coastal species . 38. P. parahypotropa (p. 1 98) 

81b Lobes more or less rounded, sublinear laciniae absent. Medulla K— , PD— . Inland 

species - 82 

82a (81b) Underside black in the centre, white peripherally. Medulla UV— (norlobaridone 

and/or protolichesterinic acid) - • 24. P. hababiana (p. 1 84) 

82b Underside almost entirely white. Medulla UV+ or UV— 83 

83a (82b) Medulla UV+(alectoronic acid) „ 34. P. louisianae (p. 195) 

83b Medulla UV- (protolichesterinic acid) . 72. P. sp.B(p. 226) 

84a (80b) Rhizines dimorphous, often extending to the margins 85 

84b Rhizines uniform, rarely extending to the margins 86 

85a (84a) Soralia marginal. Medulla PD+ orange, KC— (salazinic acid) or PD— , KC+ red 

(norlobaridone) 59. P. subsumpta (p. 215) 

85b Soralia laminal. Medulla PD-,KC- (fatty acids) 43. P. pilosa (p. 202) 

86a (84b) Upper cortex distinctly maculate - 87 

86b Upper cortex emaculate or faintly maculate 88 

87a (86a) Medulla UV+ (alectoronic acid, ± a-collatolic acid, ± gyrophoric acid) 

32. P. lobulascens (p. 192) 
87b Medulla UV— (gyrophoric acid and/or norlobaridone, ± norstictic acid) 

58. P. subschimperi (p. 2 14) 

88a (86b) Plant saxicolous 89 

88b Plant normally corticolous 91 

89a (88a) Medulla UV+ (alectoronic acid) 45. P. poolii (p. 203) 

89b Medulla UV- .90 

90a (89b) Medulla PD- (protolichesterinic acid) 23. P. grayana(p. 184) 

90b Medulla PD+ orange-red (fumarprotocetraric and protocetraric acids) 

48. P. pseudograyana (p. 206) 

91a (88b) Medulla PD+ orange to red 92 

91b Medulla PD— (but soralia may be PD+ sulphur yellow) .96 

92a (9 1 a) Protocetraric acid present 93 

92b Protocetraric acid absent 95 

93a (92a) Medulla C+ rose. Gyrophoric acid present 67. P. umbrosa (p. 222) 

93b Medulla C-. Gyrophoric acid absent 94 

94a (93b) Cilia well developed. Medulla UV+ (alectoronic or #-collatolic acid) or UV — 

(protolichesterinic acid) 55. P. subarnoldii (p. 212) 

94b Cilia poorly developed, present only in the lobe axils. Medulla UV— (± undetermined 

fatty acids) 22. P. gardneri (p. 183) 

95a (92b) Stictic acid present 40. P. perlata (p. 200) 

95b Salazinic acid present , 12. P. cristifera (p. 173) 

96a (9 lb) Soralia PD+ sulphur yellow (psoromic acid) . .... 16. P. direagens(p. 178) 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 165 

96b Soralia PD— or at most PD+ pale yellow (psoromic acid absent) 97 

97a (96b) Medulla UV+(alectoronic acid) 45 P poolii (p 203) 

97b Medulla UV- " ... 98 

98a (97b) Medulla C— (norlobaridone and/or protolichesterinic acid) 24. P. hababiana (p 1 84) 
98b Medulla C+ red 99 

99a (98b) Lecanoric acid present . .10. P. cooperi (p. 172) 

99b Gyrophoric acid present 100 

1 00a (99b) Soralia often ciliate, soredia granular. Fatty acids present . . 33. P. lophogena (p. 1 94) 
100b Soralia eciliate, soredia farinose. Fatty acids absent .101 

101a (100b) Norlobaridone present m 27. P. indoafra(p. 187) 

101b Norlobaridone absent 52. P. sancti-angelii (p. 210) 

Descriptions of the species 

1. Parmelia abessinica Nyl. ex Krempelh. 

Linnaea 41 : 140 {\%ll).—Parmotrema abessinicum (Nyl. ex Krempelh.) Hale, Phytologia 28 : 334 
(1974). Type: Abessinia, leg. Hildebrandt (G-holotype). [TLC (Winnem 1974) : norlobaridone, 
loxodin, protolichesterinic acid, atranorin.] 

Thallus corticolous, more or less coriaceous, adnate to loosely attached, pale grey. Lobes 
0-5-1 cm broad, margins ascending, crenate, ciliate, cilia 0-5-1-5 (2) mm long. Upper side 
faintly to distinctly maculate, rugose and cracked with age. Medulla white. Underside 
rugose, sparsely rhizinate, black in the centre, with a brown, mottled, or white marginal 
zone. Soralia and isidia absent. Apothecia common, often crowded, stipitate, up to 1 cm in 
diameter, thalline margin smooth to crenulate, eciliate (or very rarely ciliate), disc perforate, 
spores 15-18 (20) x 8-1 jum. Conidia weakly sublageniform, 6-8 jum long. 

TLC: (1) norlobaridone, ± loxodin, protolichesterinic acid, atranorin, (2) norlobaridone, 
± loxodin, atranorin, (3) protolichesterinic acid, atranorin, (4) fatty acids of the reddenda 
type, atranorin. 

Two specimens, 3K 23/124 and Nordal INB 820, from Kenya and Tanzania respectively, 
belong to a hitherto undescribed chemotype. They have the same general habit as P. 
abessinica, spores in the same size range, and similar conidia, and differ only in producing 
unnamed fatty acids of the reddenda type in the medulla. An additional specimen of this 
chemotype has been seen from Zaire (LD). The same chemical properties are found in P. 
mesotropa Mull. Arg., described from Paraguay, but that species lacks cilia and has 
imperforate apothecia. 

Parmelia glaucocarpoides Zahlbr., based on P. glaucocarpa Mull. Arg., non Ach., and 
described on material from Madagascar (G — holotype), was regarded by Hale (1965) as 
synonymous with P. abessinica. Its morphology is similar, it has sublageniform conidia 
5-6 jum long, and contains protolichesterinic acid, but the spores are 25-27 (30) x 12-15 jum 
and fairly thick-walled, showing that P. glaucocarpoides should be regarded as a species 
distinct from P. abessinica. 

Parmelia abessinica is a common and widespread species of dry, well lit sites at inter- 
mediate altitudes, such as bushed grassland, artificial habitats, open hillsides, and the edge of 
forests. It has been collected between 900 and 2600 m, but is most common below 2000 m. 
Outside our area it is known from South and West Africa and Mexico (Hale, 1 965). 

Selected East African records 

Ethiopia. Sidamo Province, Yirga Alem, Winnem 477/1 (O). Kenya. Eastern Province, Marsabit 
District, Mt Marsabit, vicinity of Lake Paradise, 4K 6/124: Kitui District, Mutomo, 50 km NE of 
Kibwezi, K 21/2; Machakos District, Ol Doinyo Sapuk, 20 km SE of Thika, 2K 1/101, lava How 5 km 



66 



H. KROG & T. D. V. SWINSCOW 



NW of Kibwezi, 2K22/101, 5K2/31; Embu District, Izaak Walton Inn, K 53/8; Meru District, 
between Chogoria and South Mara River, 3K 1 1/104. Rift Valley Province, Kajiado District, Chyulu 
Hills, K 39/13; Samburu District, WSW slope of Warges, 4K 3/101; Laikipia District, Burguret, 
4K 25/108; Nakuru District, Eastern Mau Forest 8 km ENE of Mau Narok, 4K 32/109, Lake Naivasha 
Hotel, on east shore of lake, 3K 21/14. Coast Province, Taita District, near school W of Wundanyi, 
2K 27/101. Western Province, Kakamega District, Kakamega Forest by River Ikuyawa, 4K 10/101. 
Tanzania. Lake Province, Mosuma District, near Fort Ikomo Lodge, Sipman 6357 (herb. Sipman). 
Northern Province, Arusha District, Arusha National Park, Juniper Hill, T 3/1 14, Mt Meru Crater, 
T 5/162; Mbulu District, Babati, Sitari 541 (TUR). Southern Highlands Province, Njombe District, 
12 km SE of Njombe, Nordal INB 820 (O). Uganda. Kigezi District, Bufumbira County, Kisoro, 
Travellers' Rest Hotel, U 19/3, 3U 59/10; Ruzhumbura County, W edge of Maramagambo Forest, 
2U 1 1/6. Karamoja District, Matheniko County, near Sogolomon on Mt Moroto, 2U 36/53, NW of Mt 
Moroto, 4 km SW of Nakiloro, 2U 38/6. 



2. Parmelia aldabrensis Dodge. 



Fig. 2 



Ann. Mo. bot. Gdn 46: 160 (1959). — Parmotrema aldabrense (Dodge) Hale, Phytologia 28: 334 
(1974). Type: Aldabra Group, coll. Walter Fox 220 (BM— holotype). [TLC (Winnem 1974): 
norstictic acid.] 

Thallus corticolous, adnate, pale grey to grey-green. Lobes 0'5-0'8 cm broad, deeply 
divided, margins ascending, crenate, ciliate, cilia up to 2*5 mm long. Upper side distinctly 
maculate. Medulla white. Underside smooth or somewhat rugose, medium to dark brown, 
rarely with a white mottled marginal zone, rhizines slender, scattered almost to the margins. 
Soralia and isidia absent. Apothecia numerous, marginal and submarginal, often 
eccentrically orientated on semitubular stipes, disc 0*5—1 (1*5) cm in diameter, with a small 
perforation, thalline margin smooth or weakly crenate, occasionally ciliate, spores 
1 3-1 7 x 5-8 jum. Conidia filiform, 12-15 jum long. 
TLC: norstictic acid, stictic acid. 

All East African specimens contained stictic and norstictic acids, but the type specimen, as 
well as two other specimens from the Aldabra Islands, Stoddart 992 (BM) and Fosberg 493 19 
(S), had only norstictic acid. Morphologically the two populations are the same. No 
apothecia or conidia were found in the holotype, but the collection Stoddart 992 had spores 




Parmelia aldabrensis Dodge, 3K 30/1 10 (O). Rule = 1 cm. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



167 



12-16 x 6-7 jum and filiform conidia 10-12 /zm, which agrees well with the East African 
material. 

Parmelia preperforata W. Culb., described from Texas (US — holotype), is chemically 
concordant with the East African specimens of P. aldabrensis, has a maculate upper cortex, 
marginal to submarginal apothecia, and similar spores and conidia. It differs in having 
broadly rounded lobes which are not particularly deeply divided, and an underside which is 
black in the centre with a wide, pale ivory marginal zone. 

Parmelia aldabrensis is locally abundant in mangroves by the Indian Ocean, where it 
often grows convoluted around thinner branches. It has also been collected in the lowlands of 
Tanzania up to 1000 m. Outside our area it is known from the Aldabra Islands and 
Madagascar (Hale, 1965). 

East African records 

Kenya. Coast Province, Kwale District, 2 km N ofGazi, K44/2, 111, 3K 30/45, 110; Kilifi District, 
Mida Creek, 3K 27/104, 3K 29/106, Gedi ruins, 3K 25/104. Tanzania. [? Tanga Province] 
Masaihochland, Wadiboma, Fischer 703 (G, type of P. hildebrandtii var. ciliata Mull. Arg.). Eastern 
Province, Mzizima [Uzaramo] District, Kunduchi, Bjornstad AB 1942 (O); Morogoro District, Mindu 
Forest Reserve, 4 May 1978, Dahl(Q). 



3. Parmelia amaniensis Steiner & Zahlbr. 



Fig. 3 

July 



Bot. Jb. 60:526 (1926). [Tanzania] Im Regenwald bei Amani, Ost-Usambara, 800 m, 
1909, leg. Brunnthaler (W — holotype). [TLC: alectoronic acid, protocetraric acid, atranorin.] 
Parmelia pachyspora Hale, Contr. U.S. natn. Herb. 36 (5) : 299 (1965). — Parmotrema pachysporum 
(Hale) Hale, Phytologia 28:338 (1974). Type: Angola, Huila, 10 km N of Sa de Bandeira, 3 
February 1960, leg. G. Degelius (herb. Degelius — holotype). [TLC: a-collatolic acid, alectoronic acid 
(+), protocetraric acid, atranorin.] 

Thallus corticolous, membranaceous to more or less coriaceous, loosely attached, pale grey. 
Lobes 0*5-1-5 cm broad, crenate, ciliate, cilia 3-5 (7) mm long. Upper side emaculate. 




Fig. 3 Parmelia amaniensis Steiner & Zahlbr., Ryvarden 1 1 632 (O). Rule = 1 cm. 



168 H. KROG & T. D. V. SWINSCOW 

Medulla white. Underside black, with a narrow brown or rarely white mottled marginal 
zone, rhizines sparse. Soralia and isidia absent. Apothecia up to 1*7 cm in diameter, 
submarginal on swollen stipes which become somewhat constricted with age, or marginal 
and eccentrically situated on semitubular, convoluted lobes, thaline margin normally 
lobulate-dentate and ciliate, disc imperforate, spores thick-walled, 25^0 x 15-18 jum. 
Conidia weakly sublageniform, 5-8 jum long. 

TLC: (1) alectoronic acid, protocetraric acid, atranorin, (2) #-collatolic acid, 
± alectoronic acid (trace), protocetraric acid, atranorin, (3) ± protolichesterinic acid, 
protocetraric acid, atranorin. 

Overlooking the presence of protocetraric acid, Hale (1965) reduced P. amaniensis to 
synonymy with P. subrugata Krempelh., a South American species with alectoronic and 
tf-collatolic acids. In the same work he described the new species P. pachyspora, with 
protocetraric acid as its diagnostic medullary substance. Later, under the description of the 
new species P. paradoxa (Hale, 1973), he noted that P. pachyspora was now known to 
contain alectoronic acid in addition to protocetraric acid. When subjecting the type speci- 
mens to TLC we found that P. amaniensis contained alectoronic acid and P. pachyspora 
mainly a-collatolic acid in addition to protocetraric acid. Since anatomy, morphology, and 
ecological requirements are the same for the two species, we here reduce P. pachyspora to 
synonymy with P. amaniensis. We also include in the species some specimens which 
produced protocetraric acid ± protolichesterinic acid in the medulla (Table 12). 

In our area the typical strain has been collected in Tanzania, the pachyspora strain in 
Uganda, and strain (3) in both countries. 

Parmelia amaniensis has been found in the lower montane forests between 800 and 
1 150 m altitude, with the exception of one collection from 2000 m. In East Africa we have 
seen specimens from Tanzania and Uganda. Outside our area the species occurs in Angola, 
Malawi, Mozambique, Sierra Leone, Zambia and Zimbabwe. It seems to be restricted to 
Africa. 

East African records 

Tanzania. Eastern Province, Morogoro District, Nguru Mountains, E slope above Kwamanga village 
near Mhonda Mission, Pocs & Mabberley 6397/H (herb. Pocs), Uluguru Mountains, N slope of 
Bondwa above Morogoro, Pocs 6732 (herb. Pocs). Tanga Province, Tanga District, East Usambara 
Mountains, Amani Forest Reserve, T. & S. Pocs 6100/AD-B (herb. Pocs); Lushoto District, in rain 
forest by Amani, East Usambara, Brunnthaler 7/1909 (W — holotype of P. amaniensis), Usambara Mts, 
Amani, Karimi Estate (road towards Monga), Moberg 1485b (UPS). Uganda. Masaka District, Bukoto 
County, Jubiya Forest, 3U 28/18, 3U 32/3, Lye L 604A (herb. Lye), N edge of Malabigambo Forest, 
3U25/4. 

4. Parmelia andina Mull. Arg. 

Revue mycol. 1 : 169 (1879). — Parmotrema andinum (Mull. Arg.) Hale, Phytologia 28 : 334 (1974). 
Type: prope Cisne, Ecuador, 2800 m, leg. Ed. Andre 4324 bis (G — holotype, BM — isotype). [TLC 
(Winnem 1975): lecanoric acid, atranorin.] 
Parmelia modesta Hue, Bull. Soc. Bot. France 63, Mem. 6 (28) : 6 (1916). Type: Afrique equatoriale 
anglaise, 1912, leg. Vte de Poncins, (PC — holotype). [TLC: lecanoric acid, atranorin.] 

Thallus corticolous or rarely saxicolous, coriaceous, loosely attached, pale grey, medium 
grey, or grey-green. Lobes up to 2 cm broad, rounded, entire or crenate, eciliate. Upper side 
usually matt, faintly to distinctly maculate (especially in the vicinity of the apothecia), 
pitted, folded, or rugose towards the centre. Medulla white. Underside black, matt and 
rugose, with a white, brown, or mottled marginal zone, rhizines short, black in central parts, 
pale brown or white peripherally, unevenly distributed. Soralia and isidia absent. Apothecia 
submarginal to laminal, often numerous, sometimes crowded and obscuring the thallus, 
1-3 (4) cm in diameter, thalline exciple rugose, strongly maculate, disc widely perforate, 
spores 13-18 x 8-10 //m. Conidia filiform, 10-16 (20) jum long. 
TLC: lecanoric acid, atranorin. 



PARMEL1A (AMPHIGYMNIA) IN EAST AFRICA 169 

Parmelia andina differs from P. hololoba mainly in the absence of cilia. When P. hololoba 
extends inland and occurs in the same habitats as P. andina, its cilia sometimes become 
sparse, short, and stunted, so that the two species are almost indistinguishable. 

Parmelia andina is common and widespread in dry, well lit sites, such as savannas, open 
hillsides, and artificial habitats. It has been collected between 900 and 2400 m altitude in all 
four countries of our studies. Outside our area it is known from numerous localities in 
continental Africa as well as from Madagascar, India, Thailand, Tahiti, and South America 
(Hale, 1965). 

Selected East African records 

Ethiopia. See Winnem (1975). Kenya. Eastern Province, Isiolo District, 8 km S of Isiolo, 4K 28/101; 
Machakos District, Kilima Kiu, 5K3/2; Embu District, Embu, Izaak Walton Inn, K 53/109; Meru 
District, vicinity of Chogoria, 3 K 9/104; Marsabit District, c. 1 km N of the mountain Ajaumarka, 
Moberg 3864 (UPS). Rift Valley Province, Elgeyo Marakwet District, 8 km NE of Kapcherop, 
2K 15/1 14; Nakuru District, E of road by Lake Elmenteita, 2K 20/101; Samburu District, WSW slope 
of Warges, 4K 3/106. Western Province, Kakamega District, Kakamega Forest by River Ikuyawa, 
4K 10/105. Nyanza Province, Kisii District, 1 km SE of Kisii, 4K 12/114. Central Province, Nyeri 
District, Mt Kenya Safari Club, 4K 19/112. Tanzania. Northern Province, Arusha District, Arusha 
National Park, Kusare Forest, T 2/120. Eastern Province, Morogoro District, above Morningside in 
the Uluguru Mts, 6 May 1978, Dahl(0). Southern Highlands Province, Iringa District, Ruaha National 
Park, summit of Magangwe Hill. Bjornstad 1780-a (O). Southern Province, Songea District, Gumbiro, 
Nordal INB 849 (O). Uganda. W. Mengo District, Kyadondo County, Makerere University Campus, 
U l/7b, Manum 1/1968 (O); Busiro County, Entebbe Botanical Garden, U 9/56b. Toro District, 
Burahya County, Fort Portal, Dale 144 (BM — type of P. dalei Dodge); Busongora County, 10 km NW 
of Kilembe, 2U 12/22. Karamoja District, Matheniko County, south side of Mt Moroto, 2U 35/4. West 
Nile District, Mt Otse, 1962, Thomas (BM — type of P. thomasii Dodge). 

5. Parmelia aprica Krog & Swinscow sp. nov. Fig. 4 

Thallus corticola, coriaceus, adnatus, flavus ad flavo-viridis. Lobi 0*5-1 cm lati, eciliati, marginibus 




Fig. 4 Parmelia aprica Krog & Swinscow, holotype (O). Rule = 1 cm. 



170 H. KROG & T. D. V. SWINSCOW 

integris vel leniter crenatis, superne emaculati, rugescentes, rimosi, Soralia marginalia, ad lobos 
peripherales linearia, ad lobos ascendentes laterales subcapitata, aliquando ad laminam extensa. 
Apothecia et pycnidia ignota. Acidum protocetraricum et acidum usnicum continens. 

Thallus corticolous, coriaceous, adnate, bright yellow to yellow-green. Lobes 05—1 cm 
broad, eciliate, with entire or weakly crenate margins. Upper side emaculate, becoming 
rugose, irregularly cracked and sometimes lobulate towards the centre. Medulla white. 
Underside black in the centre, with a broad, brown, shiny marginal zone, rhizines 
sparsely developed. Soralia marginal, linear on peripheral lobes and subcapitate on 
ascending lateral lobes, sometimes spreading diffusely on to the lamina. Soredia granular, 
pale green to yellowish. Apothecia and pycnidia unknown. 

TLC: protocetraric acid, ± traces of fatty acids, usnic acid. 

Type: Kenya. Eastern Province, Machakos District, lava flow 5 km NW of Kibwezi, 
2°35 , S, 37°51'E, 1000 m altitude, on shrubs, January 1972, coll. H. Krog & T. D. V. 
Swinscow no. K 20/103 (O— holotype; BM, UPS— isotypes). 

Parmelia aprica differs from both P. rava and P. dilatata in its bright yellow colour, yellow- 
green soredia, and the absence of atranorin in the cortex; it differs from P. dilatata also in the 
absence of echinocarpic acid and associated substances. 

Parmelia vivida, newly described in this work, appears to be the corresponding primary 
species. It has spores 20-22 x 8-10 jum and sublageniform conidia 6-7 jum long. 

Parmelia aprica is known only from the type locality, where it grew on shrubs in a rather 
dry site exposed to high light intensity. Further collection numbers are 2K 22/105 and 
3K23/1,129. 

6. Parmelia araucariarum Zahlbr. 

Denkschr. Akad. Wiss., Wien, math.-nat. Kl. 83 : 179 (1909). — Parmotrema araucariarum (Zahlbr.) 
Hale, Phytologia 28 : 334 (1974). Type: Brasilia. Prov. Sao Paulo. Prope S. Amaro in circuitu urbis 
S. Paulo, 800 m, May 1901 , leg. V. Schiffner (W— holotype, O— isotype). [TLC: pigment, fatty acids 
in the reddenda complex, atranorin.] 

Thallus corticolous, membranaceous, loosely attached, green-grey. Lobes 0*5-1 -5 cm broad, 
ciliate, cilia 1-2 (3) mm long. Upper side emaculate, more or less irregularly cracked in older 
parts. Medulla distinctly pigmented salmon pink, yellow, or ochraceous, pigment K — . 
Underside black, with a brown, naked marginal zone, rhizines sparse. Soralia marginal, 
linear. Mature apothecia not seen, but the holotype specimen has one immature, 
imperforate apothecium with a sorediate thalline margin. Conidia sublageniform, 6-8 jum 
long. 
TLC: pigment, ± protolichesterinic acid, atranorin. 

Parmelia araucariarum was described as an eciliate species by both Zahlbruckner (1909) 
and Hale (1965). Later, Hale (1974a) noted that cilia might be very sparsely developed. In 
fact, short cilia are present in an isotype in O, and the holotype, on close examination, also 
possesses one or two elusive cilia. The East African specimens are distinctly ciliate with most 
cilia less than 2 mm but occasionally reaching 3 mm in length. The species should be 
regarded as normally ciliate, and does probably not represent the sorediate counterpart of P. 
myelochroa Hale as suggested by Hale (1965). 

The type specimens contain different fatty acids from those of the East African specimens. 
We regard this as a minor chemical variation of no taxonomic consequence. 

Parmelia araucariarum is rare in East Africa. We have collected it twice in Kenya where it 
was growing in the lower montane forest on the south and east slopes of Mt Kenya at 
2000-2 1 00 m. The species was previously known only from the type locality in Brazil. 

East African records 

Kenya. Central Province, Kirinyaga District, Mt Kenya, 2 km NW of Irangi Forest Station, moist 
deciduous forest near River Ena, K 48/1 1 1 . Eastern Province, Meru District, Mt Kenya, open forest on 
east side, at Them we, 3 K 16/104, 108. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 171 

7. Parmelia austrosinensis Zahlbr. 

Symb. sin 3 : 192 (1930).— Parmotrema austrosinense (Zahlbr.) Hale, Phytologia 28 : 335 (1974). 
Type: China, Kveitschou, Gwanyinschan, near Kweiyang, leg. Handel- Mazetti 10580 (WU — not 
seen). 

Thallus corticolous or more rarely saxicolous, usually somewhat coriaceous, loosely 
attached, pale grey, ash grey, or grey-green. Lobes 1-3 cm broad, rounded, entire or crenate, 
eciliate, margins often ascending. Upper cortex weakly maculate, sometimes distinctly 
maculate, especially in central parts, more or less rugose and reticulately cracked. Medulla 
white. Underside black, with a white, brown, or mottled marginal zone, rhizines short, 
unevenly distributed. Soralia marginal and submarginal, soredia granular. Apothecia 
substipitate, thalline exciple sorediate, disc perforate or imperforate, spores 12-17 (20) x 
7-10 jum. Conidia filiform, 10-14 //m. 
TLC: lecanoric acid, atranorin. 

For differences from P. defecta, see under that species. From P. cooperi it differs mainly in 
the absence of marginal cilia. 

Parmelia austrosinensis is one of the commonest Amphigymnias in dry, well lit sites in all 
four countries of our study. It has been collected between 1000 and 3000 m altitude, but is 
most common below 2500 m. Outside our area it is a common and widespread species in 
tropical amd temperate regions. 

Selected East African records 

Ethiopia. See Winnem (1975). Kenya. Central Province, Kiambu District, escarpment E of Rift Valley, 
35 km NW of Nairobi, K 15/121; Nyeri District, Mt Kenya, W side, Naro Moru track, K 32/120; 
Kirinyaga District, Mt Kenya, S side, by Thiba Fish Camp, K 52/1 13. Rift Valley Province, Kajiado 
District, Chyulu Hills, K 39/104; Elgeyo Marakwet District, 7 km N of Chebiemit, 2K 6/127; Uasin 
Gishu District, near Sergoi, 2K 9/1 12; Trans Nzoia District, by road E of Moiben, near Hoey's Bridge, 
2K 12/1 16; Nakuru District, E of road by Lake Elmenteita, 2K 20/1 19; Samburu District, WSW slope 
of Warges, 4K 3/107; Kericho District, Cheboswa ENE of Kericho, 4K 17/113; Laikipia District, 
Burguret, 4K25/113; Narok District, Enabilibil, 22km S of Siape Bridge, 4K31/105. Eastern 
Province, Embu District, 10 km N of Embu, by River Rubingazi, K 46/106; Machakos District, Ol 
Doinyo Sapuk, 20 km SE of Thika, 2K 1/115; Meru District, S side of Chogoria, 3K 9/106. Coast 
Province, Taita District, above Wundanyi, 2K 25/133. Western Province, Kakamega District, 
Kakamega Forest by River Ikuyawa, 4K 10/103. Nyanza Province, Kisii District, 6 km E of Keroka, 
4K 11/109; Homa Bay District, 7 km WSW of Ringa, 4K 14/102. Tanzania. Northern Province, 
Arusha District, Arusha National Park, Kusare Forest, T 2/1 16. Eastern Province, Morogoro District, 
above Morningside in the Uluguru Mts, 6 May 1978, Dahl (O). Southern Highlands Province, Njombe 
District, between Kitulo and Matamba, 9 May 1978, Dahl{0). Uganda. Karamoja District, Matheniko 
County, southern slopes of Mt Moroto, 2U 35/9; Dodoth County, Kidepo Valley National Park, 
3U 15/112. W. Mengo District, Busiro County, Entebbe Botanical Garden, 3U 5/20. Kigezi District, 
Bufumbira County, 2 km W of Kanaba Gap, 3U 58/7. 

8. Parmelia bangii Vainio 

in Schmidt, Bot. Tidsskr. 29 : 104 (1909).— Parmotrema bangii (Vainio) Hale, Phvtologia 28 : 335 
(1974). Type: Lichens of South America No. 13, Bolivia, La Paz, 1891, M. £<2«g(H-NYL 35500— 
holotype). [TLC: stictic acid, constictic acid, atranorin.] 

Thallus corticolous, membranaceous, loosely attached, pale green-grey. Lobes 0*5—1 cm 
broad, crenate, sparingly ciliate, cilia 0*5-1 (2) mm long. Upper cortex emaculate, fragile, 
cracking and flaking, folded and rugose towards the centre. Medulla white. Underside 
smooth, shiny, black, with a pale brown marginal zone, rhizines mostly dense, in some 
places reaching almost to the margins. Soralia developing as pustules in a broad submarginal 
zone, soredia granular. Apothecia not seen in East African material. (In the holotype 
apothecia are subsessile, thalline exciple sorediate, disc imperforate, spores 30^0 x 
1 5-20 //m, thick- walled.) Conidia rod-shaped to shortly filiform, 8-10 //m long. 
TLC: stictic acid, constictic acid, atranorin. 



172 H. KROG & T. D. V. SWINSCOW 

Parmelia bangii has been collected in inselbergs and montane forests in Kenya between 
1 500 and 3200 m altitude. It was previously known from two localities in Bolivia and 
Colombia respectively (Hale, 1965) and from Tenerife, the Canary Islands (0sthagen & 
Krog, 1976). 

East African records 

Kenya. Eastern Province, Fort Hall District, Aberdare Mountains, Kimakia Forest Station, Ryvarden 
9028 c (O); Machakos District, Ol Doinyo Sapuk, 20 km SE of Thika, 2K 4/101 ; Marsabit District, Mt 
Marsabit, W side of Sokorte Dika 'swamp-lake', Lye L661 (herb. Lye); Meru District, Mt Kenya, N 
slope, Sirimon track, 4K 20/127. Central Province, Embu District, Embu, by Izaak Walton Inn, 
3K6/130. Rift Valley Province, Kajiado District, Ngong Hills, K 45/118. Coast Province, Taita 
District, near school W of Wundanyi, 2K26/101, 2K27/102; Taita District, Taita Hills, Mt Iyale, 
Nordal INB 725 (O). 

9. Parmelia cetrata Ach. 

Syn. Lick.: 198 (\8\4).—Parmotrema cetratum (Ach.) Hale, Phytologia 28 : 335 (1974). Type: Ad 
corticem arborum in America septentr. (not seen). 

Thallus corticolous, membranaceous to somewhat coriaceous, medium grey to green-grey. 
Lobes up to 15 cm broad, rounded, crenate or deeply incised, the laciniae 1-3 mm broad, 
with truncate apices, ciliate, cilia 1-2 mm long. Upper cortex reticulately maculate and 
cracked. Medulla white. Underside smooth, black, with a dark brown marginal zone, densely 
rhizinate, often with rhizines and papillae to the margins, rhizines mainly simple, but 
occasionally squarrose. Soralia and isidia absent. Apothecia stipitate, thalline exciple entire, 
disc perforate, spores 1 1-13 x 8-10 jum. Conidia filiform, 10-14 jum long. 
TLC: salazinic acid, atranorin (+). 

Parmelia cetrata, a widespread temperate species, is rather rare in East Africa. It has been 
collected in inselbergs and upland habitats which are generally well lit but periodically 
influenced by mist. We have found it in Kenya and Tanzania between 1400 and 2000 m 
altitude, with the exception of one collection from 2600 m. 

East African records 

Kenya. Eastern Province, Marsabit District, Mt Marsabit, vicinity of Lake Paradise, 4K 6/131; 
Machakos District, Ol Doinyo Sapuk, 20 km SE of Thika, 2K 1/104, 2K 2/125, Mua Hills, 3K 2/120. 
Coast Province, Taita District, near school W of Wundanyi, 2K 26/102, 2K 27/105. Tanzania. 
Northern Province, Arusha District, Arusha National Park, Juniper Hill, T 3/1 13, Mt Meru Crater, 
T 5/1 17. 

10. Parmelia cooperi Steiner & Zahlbr. 

in Zahlbruckner, Bot. Jb. 60 : 528 (1926). Type: South Africa, Kapland, Kapprana, Cooper 1813 
(W — holotype, not seen). 

Thallus corticolous or rarely saxicolous, membranaceous to coriaceous, loosely attached, 
pale grey to grey-green. Lobes more or less revolute, 0*8-2 cm broad, rounded, entire or 
crenate, ciliate, cilia coarse or slender, (0*3) 2-5 mm long. Upper side emaculate or faintly 
maculate, more rarely distinctly maculate, reticulately cracked towards the centre. Medulla 
white. Underside black to the margins or with a brown or mottled marginal zone, rhizines 
short, sparse, unevenly distributed. Soralia marginal and submarginal, soredia granular. Only 
immature apothecia seen in the East African material. (According to the protologue the 
spores are 15-19 x 7-9 (10)//m.) Conidia shortly filiform, 10-12 (14)//m long. 
TLC: lecanoric acid, atranorin. 

Parmelia cooperi is believed to be the sorediate counterpart of P. hololoba. They seem to 
have about the same ecological requirements, preferring dry, well lit sites at low and 
intermediate altitudes. However, P. cooperi has not been found on the coast, while it extends 
to higher elevations than P. hololoba. It has been collected between 1 500 and 3000 m, but is 
most common below 2400 m. The species is known from continental Africa south of Sahara, 
Madagascar, and India. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 173 

Selected East African records 

Ethiopia. See Winnem (1975). Kenya. Central Province, Kiambu District, escarpment E of Rift Valley, 
35 km NW of Nairobi, K 15/3, 103; Nyeri District, Mt Kenya Safari Club, 4K 19/114; Kirinyaga 
District, Irangi Forest Station, 5K4/9a. Eastern Province, Machakos District, Mua Hills, 3K2/110; 
Meru District, Mt Kenya, E side, at Themwe, 3K 16/191; Marsabit District, Mt Marsabit, NW side of 
Lake Paradise, Lye L 657A (herb. Lye). Rift Valley Province, Nakuru District, Londiani Forest SE of 
Mau Summit, 2 K 5/103; Elgeyo Marakwet District, 7 km N of Chebiemit, 2 K 6/105; Uasin Gishu 
District, 5 km NW of Timboroa Summit, 2K 19/129; Laikipia District, Naro Moru River Lodge, 
2K 34/101. Coast Province, Taita District, above Wundanyi, 2K 25/1 11. Tanzania. Northern 
Province, Arusha District, Mt Meru Crater, T 5/155. Tanga Province, Lushoto District, Usambara 
Mts, Magamba c. 4 km N of Lushoto, Ryvarden 10911 (O). Eastern Province, Morogoro District, 
Uluguru Mts, Morningside Research Station, Ryvarden 11580 (O). Southern Highlands Province, 
Mbeya District, between Kimondo and Kikondo, 9 May 1978, Dahl (O). Uganda. Toro District, 
Busongora County, 10 km NW of Kilemba, slopes of Ruwenzori, 2U 12/45; Burahya County, 1 km N 
of Fort Portal, Pentecost RE 21 (BM). Kigezi District, Kinkizi County, Mafuga Forest, 3U 39/4. 

1 1 . Parmelia crinita Ach. 

Syn. Lich.: 196 (1814). — Parmotrema crinitum (Ach.) Choicy, Bull. mens. Soc. linn. Lyon 21 : 175 
(1952). Type: North America, Muhlenberg (H — holotype). [TLC: not tested.] 

Thallus corticolous, more rarely terricolous or saxicolous, membranaceous, loosely 
attached, pale grey, ash grey, or grey-green. Lobes 0-5-1-5 (2) cm broad, crenate or 
irregularly incised, ciliate, cilia 0*5-2 (3) mm long. Upper side emaculate, smooth to faintly 
ridged or pitted, cortex fragile. Medulla white. Underside black, shiny, with a narrow, brown, 
naked marginal zone, rhizines abundant, sometimes extending to the margins. Isidia sparse 
to numerous, laminal and marginal, usually cylindrical, short, and slender but sometimes 
forming coarse, coralloid outgrowths, commonly ciliate, occasionally dissolving into soredia. 
Apothecia rare, submarginally situated on constricted stipes, thalline exciple rugose, isidiate, 
disc imperforate, spores 25-35 x 12-18 //m, thick-walled. Pycnidia not seen. 
TLC: stictic acid, constictic acid, atranorin. 

The vegetative propagules of P. crinita show a greater variation in the tropics than they do 
in the temperate region, to a certain extent recalling those of P. lophogena and P. mellissii. 
Parmelia bangii, another stictic acid producing species with large spores, is obviously very 
closely related to P. crinita as here delimited, but it is much smaller and more narrow-lobed 
and has submarginal, sorediate pustules which do not produce laminal cilia or intergrade 
with sorediate isidia. 

Parmelia crinita is one of the commoner species of inselbergs, montane forests and the 
ericaceous zone. We have collected it between 1400 and 3400 m altitude, but most 
frequently between 2000 and 3000 m. It is a common and widespread species of humid 
habitats in temperate and tropical regions. 

Selected East African records 

Ethiopia. See Winnem (1975). Kenya. Central Province, Nyeri District, Mt Kenya, W side, Naro Moru 
track, several localities, Aberdare Mts, several localities. Rift Valley Province, Kajiado District, 
Ngong Hills, K 45/105, 518; Elgeyo Marakwet District, Chebiemit Forest 1 km W of Cheptongei, 
2K 11/105; Uasin Gishu District, 5 km NW of Timboroa Summit, 2K 19/101. Eastern Province, 
Machakos District, Mua Hills, 3K 2/128; Meru District, Mt Kenya, E side, at Themwe, 3K 16/196, Mt 
Kenya, N side, Sirimon Track, 4K 20/122; Marsabit District, Mt Marsabit, vicinity of Lake Paradise, 
4K 6/132. Tanzania. Northern Province, Arusha District, Mt Meru, several localities. Eastern 
Province, Morogoro District, Kitulangala Forest Reserve, 6 May 1978, Dahl (O). Uganda. Kigezi 
District, Rubanda County, near Rwaburimbe, U 11/2; Bufumbira County, 2 km W of Kanaba Gap, 
3U 58/2b. 

12. Parmelia cristifera Taylor Fig. 5 

J. Bot., Lond. 6 : 165 (1847).— Parmotrema cristiferum (Taylor) Hale, Phytologia 28 : 335 (1974). 
Type: Calcutta, Wallich (FH — TAYL — lectotype, BM — isolectotypes). [TLC: salazinic acid, 
atranorin.] 



174 



H. KROG & T. D. V. SWINSCOW 




Fig. 5 Parmelia cristifera Taylor, K 41/107 (O). Rule = 1 cm. 



Thallus corticolous, adnate to loosely attached, pale grey. Lobes 0-5-1 cm broad, rounded, 
entire or weakly crenate, main lobes eciliate, lateral lobes and lobe axils sparingly ciliate, 
cilia 0*5-1 5 mm long, or cilia absent. Upper cortex even, emaculate. Medulla white. 
Underside smooth, shiny, black, with a brown marginal zone, rhizines short, coarse, sparse. 
Soralia mainly marginal on lateral lobes, sorediate lobes more or less ascending. Apothecia 
not found in East African material. (An isotype in BM has small, laminal apothecia with 
spores 25-30 x 1 3-1 5 jum). 
TLC: salazinic acid, atranorin. 

In the protologue for P. cristifera, Taylor (1847) cited specimens from Mauritius (Wright), 
Brazil (Gardner), and Demerara (Parker), as well as from Calcutta (W allien). We have not 
seen the Wright collection, but a specimen from Mauritius (without collector but annotated 
'type material'), in the BM, clearly corresponds to the lectotype. That is also the case with 
another specimen from Mauritius, namely the holotype of P. sieberi Dodge (FH). The 
Gardner syntype, from Brazil, is the holotype of P. gardneri, while the Parker syntype, from 
Guyana (FH), contains fatty acids and represents P. praesorediosa. 

The lectotype and two isolectotypes (BM) of P. cristifera show a species up to 30 cm in 
diameter, with peripheral lobes up to 3 cm broad, and with lateral, marginal, linear soralia 
more or less radiating towards the centre. Short, sparse marginal cilia are present in one of 
the isotypes. The East African specimens were on the whole much smaller and they were all 
sparsely ciliate. 

Parmelia stuppea Taylor is another sorediate, ciliate species with salazinic acid; it has 
been reported from, for example, Natal and Zimbabwe (Hale, 1965). Our specimen 
4K 6/133, from Mt Marsabit, may come close to that species. It had rod-shaped to shortly 
filiform conidia, 8-10 //m long, while those seen in North American material were rod- 
shaped and 5-8 //m long. However, the soralia of our specimen had a tendency to spread 
submarginally, while those of P. stuppea usually are strictly marginal. 

Parmelia hildebrandtii Krempelh., an eciliate species, was placed in synonymy with P. 
cristifera by Hale (1965), but according to the protologue it has small, subglobose spores 
11-15x1 1-12 jum. If the spore data are correct, the species must be considered to be distinct 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



175 



from P. cristifera, and should perhaps be compared with P. stuppea (spores 12-17 x 6-9 //m), 
since the presence or absence of cilia seems to be a variable character in this group. 

The determination of the East African specimens must be regarded as tentative until 
fruiting material becomes available. However, on the basis of ecological traits we doubt that 
the specimens from the coast represent P. stuppea, a largely temperate species. 

The East African specimens cannot be distinguished from P. gardneri without a chemical 
test. However, in addition to the chemical difference, P. gardneri differs from both P. 
cristifera and P. stuppea in spore size. 

Most of the specimens we have assigned to P. cristifera have been collected at about 300 m 
altitude in Kenya, where they grew on trees in the low coastal hills off the Indian Ocean, but 
we include records up to 1450 m. The species is common and widespread in the tropics and 
subtropics, but with relatively few records from Africa (Hale, 1965). 

East African records 

Kenya. Coast Province, Kwale District, Shimba Hills, 25 km SW of Mombasa, Kivumoni Forest, 
K41/102, 107, Pengo Hill, Santesson 20914 (UPS), Kwale, Forestry Station, Santesson 20802a pro 
parte (UPS). Eastern Province, Marsabit District, Mt Marsabit, vicinity of Lake Paradise, 4K 6/133. 
Tanzania. Northern Province, Moshi District, Marangu, near Kibo Hotel, Burnet 123 (BM). Uganda. 
Masaka District, Bukoto County, near Kasaka in Jubiya Forest, Lye L605 A (herb. Lye). 

13. Parmelia cryptoxantha des Abb. Fig. 6 

Mem. Inst, scient. Madagascar B, 10 : 115 (1961). — Parmotrema cryptoxanthum (des Abb.) Hale, 
Phytologia 28:335 (1974). Type: Madagascar. Foret d'Ambohitanteli (30 km NE d'Ankazobe), 
1 600 m, des Abbayes (REN — lectotype, not seen). 

Thallus corticolous, membranaceous, loosely attached, pale green-grey, lobe margins some- 
times tinged with brown. Lobes 0*8—1*2 cm broad, rounded, entire or crenate, ciliate, cilia 




Fig. 6 Parmelia cryptoxantha des Abb., 4 K 9/106(0). Rule= 1 cm. 



176 



H. KROG & T. D. V. SWINSCOW 



numerous, slender, up to 5 mm long. Upper cortex emaculate, fragile and flaking, sometimes 
with laminal cilia. Medulla with an unevenly distributed pale pink pigment. Underside 
smooth, shiny, black, with a brown marginal zone, rhizines sparse. Laminal pustules and 
open dactyls present, dissolving into corticate granules, soredia rare. Apothecia and pycnidia 
not found in East African material. (According to Hale (1965) the apothecia are imperforate 
with spores 24-28 x 10-12 jum.) 
TLC: echinocarpic acid, protolichesterinic acid, pigment, atranorin. 

Parmelia cryptoxantha is a species of shady, moist habitats in the lower montane forest. We 
have collected it between 1550 and 2100 m altitude in Kenya and Tanzania. Outside our 
area it is known from Madagascar and Natal (Hale, 1965). 

East African records 

Kenya. Central Province, Kirinyaga District, Mt Kenya, Irangi Forest Station, 5K4/7; 2 km NW of 
Irangi Forest Station near River Ena, K 48/127. Eastern Province, Meru District, Mt Kenya, E side, on 
track to Themwe, 3K 15/111; Mt Kenya, NE slope, Chigora track, Ryvarden 11640 (O). Western 
Province, Kakamega District, Kakamega Forest, 0*5 km SW of forest station, 4K 9/106. Tanzania. 
Northern Province, Arusha District, Arusha National Park, near Ngurdoto Gate, T 6/109. 



14. Parmelia defecta Hale 



Fig. 7 



Contr. U.S. natn. Herb. 36 (5) : 244 (1965).— Parmotrema defectum (Hale) Hale, Phytologia 28 : 335 
(1974). Type: Natal, District Bergville, Cathedral Peak area, Indumeni Forest, leg. O. Almborn 8934 
(LD — holotype). [TLC: lecanoric acid, atranorin.] 

Thallus saxicolous, coriaceous, adnate and strongly attached, pale grey, darker grey towards 
the centre. Lobes 0'5-L5cm broad, rounded or irregularly incised, eciliate, flat or with 
revolute margins. Upper cortex emaculate, shiny, finely cracked in older parts. Medulla 
white. Underside black, with a brown marginal zone, matt and rugose in the centre, smooth 
and shiny peripherally, rhizines sparse, fairly coarse. Soralia marginal and submarginal, with 
coarsely granular soredia. Apothecia unknown. Conidia filiform, 10-12 jum long. 
TLC: lecanoric acid, atranorin. 

Parmelia defecta has often been confused with saxicolous forms of P. austrosinensis, another 
eciliate, sorediate species with lecanoric acid. However, even when it grows on rock P. 




Fig. 7 Parmelia defecta Hale, 2 K 9/ 1 07 (O). Rule = 1 



cm 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 177 

austrosinensis has a loosely attached thallus and broader lobes with more or less ascending 
lobe margins. Parmelia defecta is believed to be the sorediate counterpart of P. soyauxii. 

We have collected P. defecta in Kenya on dry, exposed rock between 1750 and 2100 m 
altitude, often accompanied by P. pseudo gray ana. A specimen from Entebbe, Uganda, coll. 
Proctor (K, US) was cited by Hale (1965, not seen). Outside our area the species has been 
reported from South Africa and Madagascar (Hale, 1965). 

East African records 

Kenya. Eastern Province, Machakos District, Lukenia, 30 km SE of Nairobi, 3K 1/117. Rift Valley 
Province, Uasin Gishu District, near Sergoi 27 km S of Chebiemit, 2 K 9/107; Nakuru District, 
entrance of Masai Gorge, N of Lake Naivasha, K 30/104. Coast Province, Taita District, N of Mwanda 
summit, 2K 28/4, 101. 

15. Parmelia dilatata Vainio 

Acta Soc. Fauna Flora fenn. 7(7): 33 (1890). — Parmotrema dilatatum (Vainio) Hale, Phytologia 
28 : 335 (1974). Type: Sitia, Minas Gerais, Brazil, Vainio, Lichenes Brasilienses Exs. 397 (BM — 
isotypes). [TLC: protocetraric acid, echinocarpic acid, traces of pigments, usnic acid, atranorin.] 
Parmelia affluens Hale, Phytologia 22 : 141 (1971). — Parmotrema affluens (Hale) Hale, Phytologia 
28 : 334 (1974). Type: Peru, Dept. San Martin, Tingo Maria, on tree on slope E of Tingo Maria in 
jungle, 625-1 100 m, leg. H. Allard 20700 (US - holotype). [TLC: protocetraric acid, echinocarpic 
acid, pigments, undetermined substances, usnic acid, atranorin.] 

Thallus corticolous, adnate, coriaceous, yellowish grey. Lobes 1-2 cm broad, eciliate, with 
entire or crenate margins. Upper side emaculate, becoming more or less irregularly cracked 
towards the centre. Medulla white or with an obscure yellowish pink pigment. Underside 
black, with a brown, naked marginal zone, rhizines sparse. Soralia marginal and linear on 
peripheral lobes and subcapitate on ascending lateral lobes, soredia granular. (Apothecia in a 
Brazilian specimen (S) laminal, up to 1 cm in diameter, disc imperforate, thalline exciple 
sorediate, spores 25-27 x 10-12 jum.) Pycnidia not seen. 

TLC: protocetraric acid, echinocarpic acid, undetermined substances, pigments, usnic 
acid, atranorin. 

The interpretation of P. dilatata has caused many problems, mainly owing to inadequate 
knowledge of its chemical properties. Hale (1965) considered that the type material of P. 
dilatata was an abnormal collection, being eciliate and containing a trace of usnic acid. He 
placed P. robusta Degel. and P. gardneri Dodge in synonymy with it, both sparsely ciliate 
species without usnic acid. Hale (1971c) described P. affluens, with protocetraric acid, 
atranorin, and various pigments and undetermined substances, one of which has since been 
identified as echinocarpic acid. In discussing his new species Parmotrema progenes, Hale 
(1977) pointed out that this was the fertile parent morph of Parmelia dilatata, with the same 
chemistry, namely atranorin, protocetraric acid, echinocarpic acid, and associated 
unknowns. He referred specimens with atranorin and protocetraric acid only, previously 
included in P. dilatata, to P. robusta. 

One important detail in the chemistry of P. affluens and P. progenes has been largely 
overlooked, namely the presence of generous amounts of both atranorin and usnic acid in the 
cortex. As already mentioned above, Hale found usnic acid in the type specimen of P. 
dilatata. An undetermined spot of high Rf value in HEF and of a dark grey colour after 
treatment with dilute sulphuric acid and heat, reported for both P. affluens and P progenes, 
has proved to be usnic acid running more or less into other substances on the TLC plate. 

The only difference between P. dilatata and P. affluens seems to be of a chemical nature; 
morphological characters, including spore size, are the same. However, the chemical 
difference appears to be quantitative rather than qualitative. There are, for example, traces of 
pigments in an isotype of P. dilatata in the BM, and the East African specimens showed 
presence of pigments by TLC, although the medulla appeared largely white under the low 
power binocular microscope. We regard P. dilatata as a species in which the medulla is 
inherently pigmented, but with considerable variation in the concentration of both the 



178 H. KROG & T. D. V. SWINSCOW 

pigments and other undetermined medullary substances, and we therefore place P. affluens 
in synonymy with it. 

Parmelia dilatata appears to be rare in East Africa. It has been collected at one locality in 
Kenya and one in Tanzania, at 300 and 900 m altitude respectively, both within 60 km of 
the coast. 

We have seen P. dilatata, pigmented chemotype, also from West Africa: Sierra Leone, 
Kasewe Forest Reserve, Richards R7185 L (BM). It is further known from South America 
and India (Hale, 1974a). The distribution of the unpigmented (or weakly pigmented) 
chemotype is uncertain owing to nomenclatural confusion, but the species is probably 
pantropical. 

East African records 

Kenya. Coast Province, Kwale District, Shimba Hills, Makadara Forest, Santesson 20889 (UPS). 
Tanzania. Tanga Province, Usambara Mts, Amani, in the surroundings of Forestry House, Santesson 
23151 (UPS), Moberg 1472b (UPS). 

16. Parmelia direagens Hale 

Contr. U.S. natn. Herb. 36 (5) : 288 (\965).—Parmotrema direagens (Hale) Hale, Phytologia 28 : 335 
(1974). Type: South Africa, near Kimberley, leg. John Shaw {BM— holotype). [TLC (Winnem 1974): 
psoromic acid, alectoronic acid, gyrophoric acid, atranorin.] 

Thallus corticolous, loosely attached, pale grey to green-grey. Lobes 0*5-1-2 cm broad, 
rounded, entire or crenate, ciliate, cilia 1-3 mm long. Upper cortex emaculate, more or less 
cracked in older parts. Medulla white. Underside rugose, black, with a brown or rarely 
mottled marginal zone, rhizines fairly coarse, in unevenly distributed patches. Soralia 
marginal, linear. Apothecia unknown. Conidia sublageniform to rod-shaped, 7-8 //m long. 

TLC: (1) psoromic acid, gyrophoric acid, alectoronic acid, atranorin; (2) psoromic acid, 
gyrophoric acid, atranorin; (3) psoromic acid, alectoronic acid, atranorin. 

The psoromic acid is here mainly restricted to the soralia, which react PD+ sulphur yellow 
while the medulla usually is PD— . Alectoronic and gyrophoric acids are found in the 
medulla. 

Parmelia direagens has been collected on trees in well lit sites in the lower montane forest 
between 1 500 and 2600 m altitude. Outside our area it is known from South Africa (LD) and 
the Palni Hills in India (BM). 

East African records 

Ethiopia. See Winnem (1975). Kenya. Central Province, Nyeri District, Mt Kenya Safari Club, 
4K 19/126; Nyeri District, Burguret River, 4K27/107. Rift Valley Province, Laikipia District, 
Burguret, 4K 25/121. Tanzania. Northern Province, Arusha District, Mt Meru Crater, T 5/235. 
Eastern Province, Morogoro District, Uluguru Mountains, on W slope of Lupanga above Morogoro, T. 
Pocs, P. Csontos & K. Csontos 6067/A (herb. Pocs). 

17. Parmelia durumae Krog & Swinscow sp. nov. Fig. 8 

Thallus corticola, adnatus. Lobi profunde divisi, ad 1 cm lati, crenati vel irregulariter incisi, ciliis 1-3 
(6) mm longis, superne laeves, emaculati, inferne ad centrum nigri, saepe zona marginale alba. 
Soralia isidiaque nulla. Apothecia marginalia vel submarginalia, stipitata, margine thallo ciliato, 
disco vulgo imperforato. Spori 22-26 x 12-15 jum. Conidia sublageniformia ad bacilliformia, 5-7 jum 
longa. Acidum alectoronicum, acidum a-collatolicum, et atranorinum continens. 

Thallus corticolous, adnate or with more or less free lobe margins when growing on twigs, 
pale grey. Lobes deeply divided, up to 1 cm broad, crenate or irregularly incised, ciliate, 
cilia slender, 1-3 (6) mm long. Upper side smooth, emaculate except for vicinity of 
apothecia which is maculate, central parts often strongly rugose. Medulla white. Underside 
matt, rugose, black in the centre but soon turning dark brown, commonly with a wide, white 
or mottled marginal zone, rhizines short, sparse, restricted to the centre of the thallus. Soralia 
and isidia absent. Apothecia usually numerous, marginally and submarginally situated, 
stipitate, stipe sometimes formed by a convoluted lobe, thalline exciple smooth or ridged, 




Fig. 8 Parmelia durumae Krog & Swinscow, holotype (O). Rule = 1 cm. 

strongly maculate, thalline margin weakly dentate, ciliate, disc usually imperforate, 
becoming radially split with age, spores 22-26 x 12-15 jum. Pycnidia numerous, sub- 
marginal, conidia weakly sublageniform, 5-7 //m long. 

TLC: alectoronic acid, #-collatolic acid, atranorin. 

Type: Kenya. Coast Province, Kwale District, 2 km N of Gazi, 4° 22' S, 39° 30' E, at the 
edge of a mangrove, February 1974, Coll. H. Krog & T. D. V. Swinscow no. 3K 30/132, 
(O— holotype; BM, UPS— isotypes). 

There are a number of alectoronic acid producing primary species with ciliate lobe margins 
and imperforate apothecia, but none of them seems to correspond to our species. Several of 
them, such as P. argentina Krempelh., P. maclayana, P. subproboscidea Lynge, P. 
uberrima, and P. wainii A. L. Sm., have smaller spores and differ also in various other 
characters. Parmelia stenopteris Kurok. differs chemically in that it produces fatty acids 
instead of (Z-collatolic acid; it has apothecia with a strongly dentate-lobulate thalline margin 
and a rugose exciple. Parmelia subrugata has thick-walled spores more than 30 jum long. 
Parmelia pigmentifera has similar ecology as P. durumae and resembles it morphologically; 
furthermore, both species give a strong UV+ reaction in the medulla. However, P. 
pigmentifera has a brown marginal zone on the underside and produces an ochraceous 
pigment and undetermined substances in the medulla. 

Some of the Ugandan specimens have extremely long, dense cilia. They approach a 
species from Nigeria, issued as no. 3 1 in Almborn, Lichenes africani under the name P. 
subrugata Krempelh. 

Parmelia durumae grows on twigs and branches at the edge of mangroves and in low 
coastal hills off the Indian Ocean, from sea level to c. 300 m altitude, and has also been 
collected in Uganda at about 1 100 m. 

East African records 

Kenya. Coast Province, Kwale District, 2 km N of Gazi, K44/112, 3K 30/132 (type collection), 

Shimba Hills, 25 km SW of Mombasa, Kivumoni Forest, K 41/1 , 104, Tschamtei in Duruma, January 



180 H. KROG & T. D. V. SWINSCOW 

1877, Hildebrandt 2333 (M); Kilifi District, Mida Creek, 3K 29/1 13. Uganda. Busoga District, Mutai, 
December 1950, Wood (BM). Masaka District, Bukoto County, Jubiya Forest, 3U 32/107, Lye L 604 
(herb. Lye). Toro District, Bwamba, Ituri Forest, Pentecost RE 240 (BM). N Mengo District, 7 miles N 
of Nakasongola, 1918, Langdale- Brown (EA). 

18. Parmelia eciliata (Nyl.) Nyl. 

in Fournier, Mex. PL 3 : (1872). — Parmelia crinita var. eciliata Nyl., Flora, Jena 52 : 291 (1869). — 
Parmotrema eciliatum (Nyl.) Hale, Phytologia 28 : 336 (1974). Type: Mexico, Orizaba, E. Bourgeau 
s.n. (H-NYL 35295 — holotype). [TLC: stictic acid, constictic acid, atranorin.] 
Parmelia eurycarpa Steiner & Zahlbr., Bot. Jb. 60 : 530 (1926). Deutsch Ostafrica, Ost-Usambara, im 
Regenwald auf Baumstammen b. Amani, 800 m, leg. Brunnthaler (W — holotype). [TLC: stictic 
acid, constictic acid, atranorin.] 

Thallus corticolous, loosely attached, pale grey. Lobes 0-5-0*8 cm broad, crenate or 
irregularly incised, ciliate, cilia 1-2*5 mm long. Upper cortex emaculate, rugose towards the 
centre. Medulla white. Underside smooth, black, with a brown marginal zone, rhizines 
abundant. Soralia and isidia absent. Apothecia numerous, stipitate, thalline exciple more or 
less rugose, disc imperforate, spores 26-30 x 18-20//m, thick- walled. Conidia rod-shaped. 
c.8 //m long. 
TLC: stictic acid, constictic acid, atranorin. 

Parmelia eciliata is rare in East Africa. It has been collected in Tanzania in the lower 
montane forest between 800 and 1200 m altitude. According to Hale (1965) it is widespread 
in the West Indies and in Central and South America; it was reported from Japan and 
Taiwan by Kurokawa (1968) and it is also known from South Africa. 

East African records 

Tanzania. Eastern Province, Morogoro District, Mindu Forest Reserve, 4 May 1978, Dahl (O). Tanga 
Province, Tanga District, East Usambara Mountains, Amani Forest Reserve, T. & S. Pocs 6086/T 
(herb. Pocs), East Usambara by Amani, Brunnthaler ( W — holotype of P. eurycarpa). 

19. Parmelia erubescens Stirton 

Scott. Nat. 4 : 201 (1877-78). Type: Queensland, Brisbane, F. M. Bailey 1 1 (BM— holotype). [TLC 
(Winnem 1974): salazinic acid, atranorin.] 
Parmelia virens Mull. Arg., Flora, Jena 69:255 (1886). Type: Queensland, Toowoomba, 1882, 
Hartmann (G — holotype). [TLC: salazinic acid, atranorin (+).] 

Thallus somewhat coriaceous, corticolous, loosely attached, pale grey-green. Lobes 
0*5-1 cm broad, rounded, crenate, ciliate, cilia 0*8-1*5 mm long. Upper side in part 
maculate, rugose and cracked towards the centre. Underside black, with a brown marginal 
zone, rhizines dimorphous, here and there reaching almost to the margins. Soralia and isidia 
absent. Only immature apothecia and pycnidia found in the East African specimen. (The 
type of P. virens has spores 10-12 x 8-10 //m, and filiform conidia 10-12 (14) jum long.) 
TLC: salazinic acid, norlobaridone, atranorin (+). 

Hale (1965) regarded P. erubescens as conspecific with P. subcaperata Krempelh., a species 
which has proved to contain considerable amounts of usnic acid in the cortex in addition to 
atranorin. We believe that P. subcaperata is distinct from P. erubescens, and that it is closely 
related to the isidiate species Parmotrema neotropicum Kurok. 

The East African specimen was small and immature, and did not show the strongly 
maculate upper side which is so typical for this species group. However, features of the 
underside, the presence of some maculate areas, and chemical properties leave little doubt 
about its taxonomic position. 

Parmelia recipienda Nyl., with norlobaridone, is probably conspecific with P. erubescens 
(Table 1 3). For further discussion, see under 'Circumscription of the species'. 

Interpreted in its wide sense, P. erubescens is known from Australia and Brazil, and from 
one locality in Kenya where it grew in a well lit site at 1400 m altitude. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 

East African record 

Kenya. Eastern Province, Machakos District, Ol Doinyo Sapuk, 20 km SE of Thika, 2 K 1/108. 

20. Parmelia euneta Stirton 



181 



Fig. 9 

Scott. Nat. 4:298 (1877-78).— Par motre ma eunetum (Stirton) Hale, Phytologia 28:336 (1974). 

Type: Africa (tropical), Victoria, Thomson s.n. (BM — holotype). [TLC: gyrophoric acid, atranorin.] 
Parmelia composita Hale, Phytologia 23 : 243 (1972). — Parmotrema compositum (Hale) Hale, 

Phytologia 28 : 335 (1974). Type: Tanzania, Arusha Province, Mt Meru, E slope, c. 2 km N of Kitoto 

Camp, alt. 2400 m, Santesson 22985a (UPS — holotype, BM — isotype). [TLC: gyrophoric acid, 

norstictic acid, atranorin.] 
Parmelia spilota Hale, Phytologia 27:3 (1973). — Parmotrema spilotum (Hale) Hale, Phytologia 

28:339 (1974). Type: Tanzania, Horombo Hut, Kilimanjaro, 3750 m, 9 January 1969, H. D. 

Griffin (O — isotype). [TLC: norlobaridone, norstictic acid, atranorin.] 

Thallus corticolous or saxicolous, membranaceous to somewhat coriaceous, loosely 
attached, pale grey to grey-green. Lobes (0*5) 1-2 (3) cm broad, entire, crenate, or irregularly 
incised, ciliate, cilia 1-3 (4) mm long. Upper side strongly maculate, cracked in older parts, 
here and there with discoloured areas. Medulla white, with occasional patches of an 
ochraceous, K+ purple pigment (skyrin) near the lower cortex. Underside black, with a 
brown, naked marginal zone, sometimes white mottled underneath the apothecia, rhizines 
coarse or slender, often branched, sparse to abundant, unevenly distributed. Soralia and 
isidia absent. Apothecia submarginal, stipitate on more or less swollen stipes, thalline 
exciple rugose, thalline margin crenate-dentate, disc up to 2 cm in diameter, commonly 
perforate but sometimes imperforate and radially split, spores 20-26 (30) x 12-15 (17)/zm, 
relatively thick- walled. Conidia filiform, 12-16 (20) jum long. 

TLC: (1) gyrophoric acid, ± norstictic acid, atranorin; (2) norlobaridone, ± norstictic 
acid, atranorin; (3) gyrophoric acid, norlobaridone, ± norstictic acid, atranorin; (4) 
norstictic acid, atranorin (rare). 

Hale (1965) considered P. euneta to be a species with a normally ciliate thalline exciple, a 
faintly to distinctly maculate upper cortex, and gyrophoric acid in the medulla. Later, Hale 
(1972) described the new species P. composita, containing gyrophoric and norstictic acids. It, 
too, had a maculate upper cortex, and the spores were 22-24 //m, that is, in the same size 




Fig. 9 Parmelia euneta Stirton, E 30/2 1 (O). Rule = 1 cm. 



182 H. KROG & T. D. V. SWINSCOW 

range as those reported for P. euneta. Winnem (1975), who had not seen the type of P. 
euneta, found no gyrophoric acid containing specimens with a ciliate thalline exciple in her 
Ethiopian material and concluded that the species was not represented in her collections. On 
the other hand, she found no morphological differences between specimens with only 
gyrophoric acid and specimens with both gyrophoric and norstictic acids, and consequently 
included both chemotypes in P. composita, regarding the norstictic acid as an accessory 
substance. 

The type of P. euneta, in the BM, is a small scrap about 2x3 cm, but it shows the most 
important characters of the species, such as marginal cilia, an eciliate thalline exciple, a 
strongly maculate upper cortex, fairly thick- walled spores 23-26 x 13-15//m and filiform 
conidia 12-15 pm long, agreeing fully with P. composita. We follow Winnem (1975) in 
regarding norstictic acid as an accessory substance and reduce P. composita to synonymy 
with P. euneta. 

In 1973 Hale described P. spilota, a species with norlobaridone and norstictic acid in the 
medulla and a maculate upper cortex; apothecia were unknown at the time. Winnem (1975) 
referred Ethiopian specimens with only norlobaridone to the same species, again regarding 
norstictic acid as an accessory substance. She found that the species had perforate apothecia 
and spores 20-25 pm long. In Kenya and Uganda we have found morphologically similar 
specimens with a combination of gyrophoric acid and norlobaridone, still with norstictic 
acid as an accessory substance. We consider that this chemotype connects P. spilota with P. 
euneta, and regard the two species as synonymous (Table 11). For further discussion, see 
under 'Circumscription of the species'. 

One of our specimens, from Tanzania (T 16/118), had only norstictic acid but 
corresponded to P. euneta in all other characters. It had spores 25-28 x 12-15 pm, and 
filiform conidia 12-14 jum long. 

Parmelia euneta is a mainly corticolous species of inselbergs and montane forests, 
extending into the low alpine zone where it often grows on rock. It has been collected 
between 1600 and 3750 m altitude. Outside our area the species has been reported from 
Sierra Leone, Angola, Zaire, Sri Lanka, and Haiti (Hale, 1965). However, these records need 
verification, since Hale interpreted the species in a wide sense, including, for example, 
specimens with a ciliate thalline margin of the apothecia and weakly developed maculae. 

Selected East African specimens 

Ethiopia. Presso la chiesa di Sciaura Mariam Alefa, Pichi-Sermolli 70 (BP). For further records, see 
Winnem (1975) under P. composita and P. spilota. Kenya. Rift Valley Province, Uasin Gishu District, 
5 km NW of Timboroa Summit, 2K 19/1 10; Laikipia District, Burguret, 4K 25/124; Trans Nzoia 
District, Mt Elgon crater, at Maji ya Moto (hot springs), Hedberg 949, 950 m (UPS); Kajiado District, 
Ngong Hills, K 45/18, 1 19. Central Province, Nyeri District, Burguret River, 4K 27/108, Mt Kenya, W 
side, Naro Mom track, K 34/108; Kirinyaga District, Mt Kenya, S slope, 2 km NW of Irangi Forest 
Station near River Ena, K 48/109. Eastern Province, Meru District, Mt Kenya, E side, at Themwe, 
3K 16/1 12, Mt Kenya, N side, Sirimon track, 4K 24/1 16. Tanzania. Northern Province, Arusha 
District, Mt Meru Crater, T 5/1 13; Moshi District, Kilimanjaro, Horombo Hut, Griffin (O — isotype of 
P. spilota); Masai District, Ngorongoro Crater, on the rim, near the post office at Ngorongoro, 
Sipman 6381 A (herb. Sipman). Southern Highlands Province, Mbeya District, between Kimondo and 
Kikondo, 9 May 1978, Dahl (O). Uganda. Bugisu District, N Bugisu County, by Sasa Hut on Mt Elgon, 
2U 44/62. 

21. Parmelia eurysaca Hue 

Nouv. Archs Mus. Hist, nat., Paris IV, 1 : 194 (1899). — Parmotrema eurysacum (Hue) Hale, 
Phytologia 28 : 336 (1974). Type: Mexico, 1865-66, Bourgeau (PC— holotype). [TLC: not tested.] 

Thallus corticolous, membranaceous, loosely attached, pale grey-green. Lobes 0*8-1*2 cm 
broad, rounded, entire or crenate peripherally, laciniate-lobulate towards the centre of the 
thallus, ciliate, cilia slender, simple or bifurcate, 2-3 mm long. Upper side emaculate. 
Medulla white. Underside black, with a brown, shiny, naked marginal zone, rhizines slender, 
simple or bifurcate, unevenly distributed. Soralia and isidia absent. No apothecia or pycnidia 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



183 



seen in the East African specimen. (Mexican specimens in O have stipitate apothecia with 
perforate discs, and spores 12-16 x 8-10 pirn. The conidia are rod-shaped to shortly filiform, 
8-10 jum long.) 
TLC: salazinic acid, atranorin. 

It is with considerable hesitation that we identify our sterile specimen as P. eurysaca, but the 
laciniate central lobes and the content of salazinic acid point to that species. 

The specimen grew in a rain forest in the Uluguru Mountains in Tanzania between 1 300 
and 1800 m altitude. The species has its main area in the southern United States and Mexico 
(Hale, 1965). 

East African record 

Tanzania. Eastern Province, Morogoro District, rain forest above Morningside in the Uluguru 

Mountains, 6 May 1978, Dahl{0). 



22. Parmelia gardneri Dodge 



Fig. 10 



Ann. Mo. bot. Gdn 46 : 179 (1959). Type: Brazil, Gardner (FH — holotype). [TLC: protocetraric acid, 
atranorin.] 

Thallus corticolous, coriaceous, adnate or loosely attached, pale grey to green-grey. Lobes 
0*8—1 *5 (2) cm broad, crenate, irregularly incised, or sublaciniate, eciliate or rarely sparingly 
ciliate, cilia 0'2-0*5 mm long, lobe margins more or less ascending. Upper side emaculate or 
faintly maculate. Medulla white. Underside black, with a broad, brown or mottled, naked 
marginal zone, rhizines short, sparse. Soralia marginal and submarginal, forming complex, 
raised sorediate structures on narrow lateral lobes, soredia granular. Apothecia laminal, 
subsessile, thalline exciple sorediate, disc usually imperforate, spores 18-22 (24) x 8- 
10 (1 l)//m. Conidia sublageniform, 6-7 jum long. 
TLC: protocetraric acid, ± fatty acids, atranorin. 




Fig. 10 Parmelia gardneri Dodge, K 43/ 1 1 (O). Rule = 1 cm. 



184 H. KROG & T. D. V. SWINSCOW 

We regard P. gardneri as the sorediate counterpart of P. zollingeri. It has often been 
determined as P. dilatata, but differs in the smaller spores, lack of usnic acid in the cortex, 
and a less complicated medullary chemistry. From P. robusta Degel., another sorediate 
species with protocetraric acid, it differs in a more coriaceous thallus and in ecological traits. 
(In the Canary Islands, for example, P. robusta is a species of the laurel forest at about 
800-1200 m altitude. If it occurred in our area it would most certainly grow in the upper 
montane forest. It appears to have a mainly Atlantic-Lusitanian distribution, and there are 
no reliable records of it from continental Africa.) 

Parmelia gardneri grows on twigs, branches, and tree boles, and can, when well developed, 
reach a diameter of 30 cm but is usually smaller. It grows in mangroves and in the low 
coastal hills off the Indian Ocean, and occurs farther inland at around 1000-2000 m altitude. 
All collecting sites were fairly dry and well lit. The species is believed to be more or less 
pantropical. 

East African records 

Kenya. Eastern Province, Machakos District, lava flow 5 km NW of Kibwezi, K 20/145, 3K 23/126; 
Meru District, near Chogoria, 3K7/110, 3K 9/102, Mt Kenya, E side, at Themwe, 3K 16/197. Coast 
Province, Kwale District, Kwale, Forestry Station, Santesson 20802a pro parte (UPS), Shimba Hills, 
25 km SW of Mombasa, Kivumoni Forest, K 41/101, K 42/101, Pengo Hill, Santesson 20896 (UPS), 
Makadara Forest, K43/3, 2 km N of Gazi, K44/31, 105, 3K30/33, 146; Kilifi District, Gedi ruins, 
3K 25/105, Mida Creek, edge of mangrove, 3K 29/1 18. Tanzania. Lake Province, Mwanza District, 
Massanza, Sitari 609 (TUR). Northern Province, Moshi District, by track from Marangu to Mandara 
Hut, Burnet 161 (BM). Eastern Province, Morogoro District, Kitulangala Forest Reserve, 6 May 1978, 
Dahl (O). Tanga Province, Lushoto District, East Usambara Mts, Amani Forest Reserve, T. & S. Pocs 
6086/O (herb. Pocs), Amani, surroundings of Forestry House, Moberg 1471a (UPS), Amani, road 
towards SW, Santesson 23406 (UPS). Uganda. W Mengo District, Busiro County, Entebbe Botanical 
Garden, 3U 5/1. Masaka District, Bukoto County, near Kasaka in Jubiya Forest, Lye L 606 (herb. 
Lye). 

23. Parmelia grayana Hue 

Nouv. Archs Mus. Hist, nat., Paris IV, 1 : 184 (1899). — Parmotrema grayanum (Hue) Hale, 
Phytologia 28 : 336 (1974). Type: India, in Coonoor in montibus Nilgherrensibus, 1893, leg. Gray 
(PC— holotype). [TLC: not tested.] 

Thallus saxicolous, firmly attached, pale grey. Lobes up to 0*7 cm broad, rounded, margins 
ascending, crenate, ciliate, cilia 0*5— 1*5 mm long. Upper side emaculate. Medulla white. 
Underside black, with a brown, naked marginal zone, rhizines coarse. Soralia marginal and 
submarginal, soredia often with a grey-brown tinge. Apothecia absent in East African 
material. (According to the protologue, the spores are 15-16 x 7-8 jum.) Conidia sublageni- 
form to rod-shaped, 8-10 fjtm long. 
TLC: protolichesterinic acid, atranorin. 

Parmelia grayana appears to be rare in our area; it has been collected twice in Uganda on 
exposed rock at 1 160 and 1500 m altitude. It is also known from South Africa, the Canary 
Islands, India, and Japan (Hale, 1965), Australia (Kurokawa, 1969), and Arizona (BM). 

East African records 

Uganda. Masaka District, Bukoto County, SE of Lake Kayanja, Lye L710 (herb. Lye). Ankole 

District, Igara County, south side of Lubare Ridge, 11 km SofRubirizi, 2U 15/1. 

24. Parmelia hababiana Gyelnik 

Reprium nov. Spec. Regni veg. 29:288. (1931). — Parmotrema hababianum (Gyelnik) Hale, 
Phytologia 28 : 336 (1974). Based on Parmelia abessinica var. sorediosa Mull. Arg., Flora, Jena 
68 : 501 (1885). Type: Abessinia, prope Habab, Hildebrandt 310 pp. (G — lectotype). [TLC: proto- 
lichesterinic acid, atranorin.] 

Thallus mainly corticolous, membranaceous to coriaceous, more or less adnate but with 
ascending lobe margins, pale grey to grey-green. Lobes 0*5-1 5 cm broad, ciliate, cilia 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 185 

02-25 (4) mm long. Upper cortex faintly to rarely distinctly maculate, rugose towards the 
centre. Medulla white. Underside smooth or rugose, black, with a brown, mottled, or white 
marginal zone, rhizines in scattered groups or more or less evenly distributed. Soralia 
marginal and submarginal. Apothecia with entire or sorediate thalline margin, disc 
perforate, spores 13-17 (20) x 8-10 jum. Conidia weakly sublageniform, 6-8 //m long. 

TLC: (1) norlobaridone, ± loxodin, protolichesterinic acid, atranorin; (2) norlobaridone, 
± loxodin, atranorin; (3) protolichesterinic acid, atranorin. 

In dry, well lit sites P. hababiana has a rather coriaceous, grey thallus with short marginal 
cilia, and a rugose under side with a mottled or white marginal zone. However, when it grows 
in more shady habitats, the thallus may be membranaceous, grey-green, with fairly long 
marginal cilia, and have a smooth underside with a brown marginal zone. There is no 
correlation between morphological and chemical variation. 

Parmelia hababiana is one of the commonest lichens of dry, well lit upland habitats. It has 
been collected between 800 and 2650 m altitude, but is most common between 1500 and 
2000 m. The species is particularly widespread in Africa, and is, according to Hale (1965), 
known also from India and the Americas. 

Selected East African records 

Ethiopia. See Winnem (1975). Kenya. Central Province, Kiambu District, escarpment E of Rift Valley, 
35 km NW of Nairobi, K 15/106; Nyeri District, Mt Kenya, W side, by Naro Moru track, K 33/104; 
Kirinyaga District, Mt Kenya, S side, 2 km NW of Irangi Forest Station near River Ena, K 48/1 37; 
Embu District, Embu, by Izaak Walton Inn, 3 K 6/103. Eastern Province, Machakos District, Kilima 
Kiu, 70 km SE of Nairobi, K 54/2 lb; Meru District, between Chogoria and South Mara River, 
3K 11/103. Rift Valley Province, Elgeyo Marakwet District, Sogotio Forest 8 km N of Chebiemit, 
2K 10/104; Nakuru District, cliff E of road by Lake Elmenteita, 2K 20/1 15; Kericho District, Kericho 
Tea Hotel, 4K7/110; Samburu District, WSW slope of Warges, 4K3/114. Western Province, 
Kakamega District, Kakamega Forest, 0*5 km SW of forest station, 4K9/1 13. Coast Province, Taita 
District, above Wundanyi, 2K 25/101. Tanzania. Northern Province, Arusha District, Arusha 
National Park, Juniper Hill, T 3/1 15; Mbulu District, Babati, Sitari 551-a (TUR). Uganda. Mubende 
District, Buwekula County, Mubende Hill, 3U 12/5. Kigezi District, Rubanda County, 8 km W of 
Hamurwa, 3U 47/2. Toro District, Busongora County, Kyabikere, 7 km N of Bwera, 2U 5/5. 



25. Parmelia hanningtoniana Mull. Arg. Fig. 11 

Flora, Jena 73 : 339 (1890). — Parmotrema hanningtonianum (Mull. Arg.) Hale, Phytologia 28 : 336 
(1974). Type: Trop. Afric. (or.) 2-7° lat. austr. Rev. J. Hannington (G — holotype, BM — isotype). 
[TLC: gyrophoric acid, fatty acids, atranorin.] 

Thallus corticolous, loosely attached, pale green-grey. Lobes up to 1 cm broad, ciliate, cilia 
1-3 mm long. Upper side smooth to wrinkled, distinctly maculate. Medulla white. 
Underside dark brown, strongly rugose, with scattered, agglomerated, black granules 
interspersed with groups of coarse, black rhizines. Soralia and isidia absent. Apothecia 
marginal and submarginal, urceolate, up to 15 mm in diameter, disc perforate or 
imperforate, sometimes radially split, thalline exciple covered with coarse, isidioid 
protuberances which frequently bear cilia, spores 18-20 x 8-10 //m. Conidia sublageniform, 
5-7 jum long. 
TLC: gyrophoric acid, fatty acids, atranorin. 

Hale (1965) believed that P. hanningtoniana was restricted to the savannas of West Africa, 
but the type specimen probably came from Tanzania, where Hannington collected 
extensively in central parts of the country. The specimen AB 1 389— g, cited below, grew just 
south of Hannington's area. Outside East Africa the species is known from Angola (BM, FI) 
and Zaire (Hale, 1965). There is one specimen from Brazil, coll. Richard, in the BM, and 
another specimen from the same country mixed with the type specimen of P. subcaperata 
Krempelh. in M. These are the first records of the species from South America. All known 
localities lie between the equator and c. 20° S. 



186 



H. KROG & T. D. V. SWINSCOW 




Fig. 11 Parmelia hanningtoniana Mull. Arg., Bjornstad AB 1389-g(0). Rule = 1 cm. 



East African records 

Tanzania. Trop. Afric. or., 2-7° S, Hannington s.n. (G, BM — type collection). Southern Highlands 

Province, Iringa District, 50 km S of Iringa along road to Mbeya, 1700 m alt., Bjornstad AB 1 389— g 

(O). 



26. Parmelia hololoba Hale 



Fig. 12 



Contr. U.S. natn. Herb. 36 (5) : 293 (1965). — Parmotrema hololobum (Hale) Hale, Phytologia 
28 : 336 (1974). Type: Uganda, Entebbe, 3 March 1959, Proctor (packet A) (BM— holotype). [TLC: 
lecanoric acid, atranorin.] 

Thallus corticolous, loosely attached, pale grey. Lobes up to 2 cm broad, rounded, entire or 
crenate, flat or revolute, ciliate, cilia (0*5)2-4 mm long. Upper side emaculate or faintly 
maculate. Medulla white. Underside black, with a brown or mottled marginal zone, rhizines 
short, unevenly distributed. Soralia and isidia absent. Apothecia marginal to submarginal, 
stipitate, up to 3 cm in diameter, thalline exciple smooth or rugose, strongly maculate, 
disc perforate, often eccentrically orientated in relation to the stipe, spores 
(12)15-18 (20) x (6) 8-10 /an. Conidia filiform, 12-16 /an. 
TLC: lecanoric acid, atranorin. 

Parmelia hololoba is particularly well developed in habitats close to the sea coast. It is then 
emaculate, has well developed cilia, a smooth thalline exciple, and a mainly brown marginal 
zone on the under side. In inland sites the cilia often become short and stunted, the cortex 
more or less maculate, the thalline exciple rugose, and the underside white mottled towards 
the margins, in short, it becomes indistinguishable from P. andina except for the 
presence of rudimentary cilia. We consider this morphological convergence to be environ- 
mentally induced, and rely on the presence or absence of cilia for the separation of the two 
species. 

As here delimited, P. hololoba grows at sea level by the Indian Ocean and in well lit upland 



P ARM ELI A (AMPHIGYMNIA) IN EAST AFRICA 187 

habitats more or less influenced by mist, between 1000 and 1800 m altitude. It was 
previously known only from the type locality. 

East African records 

Kenya. Coast Province, Kwale District, 2 km N ofGazi, K 44/25, 105, 3K 30/21, 115; Kilifi District, 
Mida Creek, 3K 29/122, Gedi ruins, 3K 25/106. Eastern Province, Machakos District, Ol Doinyo 
Sapuk, 20 km SE of Thika, 2K 1/103. Rift Valley Province, Kajiado District, Chyulu Hills, K 39/5A, 
103; Samburu District, WSW slope of Warges, 4K3/105; Laikipia District, Burguret, 4K25/131. 
Tanzania. Northern Province, Arusha District, Arusha National Park, Kusare Forest, T 2/126. Tanga 
Province, Lushoto District, Usambara Mts, Magamba c. 4 km N of Lushoto, Ryvarden 10912 (O), near 
Amani, T. & S. Pocs 6087/A (herb. Pocs). Eastern Province, Morogoro District, N Uluguru Mts., gorge 
of Mt Kinazi above Morogoro, Pocs 6289/AY (herb. Pocs). Uganda. W Mengo District, Busiro 
County, Entebbe, Proctor (BM — type collection). 



27. Parmelia indoafra Krog & Swinscow nom. nov. 



Fig. 13 



Parmotrema indicum Hale, Mycotaxon 5 : 436 (1977), non Parmelia indica Sprengel (1827). Type: 
India, Tamil Nadu, Kodaikanal, forest below Silver Cascade, on rocks in stream, Hale & 
Patwardhan 43874 (US — holotype, BM — isotype). [TLC: gyrophoric acid, norlobaridone, 
atranorin.] 

Thallus corticolous (or saxicolous), coriaceous, pale grey. Lobes 1-2 cm broad, crenate, 
ciliate, cilia 0'5-2 (3) mm long. Upper cortex emaculate, rugose and irregularly cracked 
towards the centre, sometimes pruinose peripherally. Medulla white. Underside smooth to 
rugose, black, with a brown or mottled marginal zone, rhizines numerous, some of them 
fairly coarse, occasionally branched. Soralia marginal on some main lobes and on numerous 
narrow, ascending lateral lobes, here and there spreading submarginally. Apothecia 
unknown. Conidia shortly filiform, 10-12 (14) jum long. 
TLC: gyrophoric acid, norlobaridone, atranorin. 




Fig. 12 Parmelia hololoba Hale, Ryvarden 1 09 1 2 (O). Rule = 1 cm. 



88 



H. KROG & T. D. V. SWINSCOW 



The species was recently described from India (Hale, 1977), and it is here reported for the 
first time from Africa. The Ethiopian specimens cited below were included in P. balensis 
(=P. subschimperi) by Winnem (1975). The two species have the same chemical properties, 
but P. subschimperi has a strongly maculate upper cortex and different ecological require- 
ments. 

Parmelia indoafra is in East Africa an upland species of fairly dry and well lit sites with 
local influence of mist. It has mainly been collected between 1450 and 1900 m altitude, with 
the exception of one collection from 2600 m. 

East African records 

Ethiopia. Sidamo Province, 10 km NW of Wadera, Winnem 595/8, 9 (O); 12 km S of Kebre Mengist, E 
13/14; Yirga Alem, Sidamo Provincial Hospital, E 20/10. Kenya. Eastern Province, Marsabit District, 
Mt Marsabit, vicinity of Lake Paradise, 4K 6/129. Tanzania. Northern Province, Arusha District, Mt 
Meru Crater, T 5/161, Arusha National Park, Kusare Forest, T 2/101, near Ngurdoto gate, T 6/108, 
111. Uganda. Toro District, Burahya County, 1 km N of Fort Portal, Pentecost RE 23 (BM). 

28. Parmelia inexpectata des Abb. 

Bull. Inst.fr. Afr. noire A, 20 : 16 (1958). — Parmotrema inexpectatum (des Abb.) Hale, Phytologia 
28 : 337 (1974). Type: Cote d'lvoire (Afr. Occ), in Monte Tonkoui, prope Man, ad truncum 
Cinchonae, 1 150 m, 16 October 1951, des Abbayes (US — isotype). [TLC: the lividic acid complex, 
atranorin.] 
Parmelia paradoxa Hale, Phytologia 27 : 1 (1973). — Parmotrema par adoxum (Hale) Hale, Phytologia 
28 : 338 (1974). Type: Uganda, Masaka District, Bukoto County, northern edge of Malabigambo 
Forest, 1 100 m, December 1971, coll. T. D. V. Swinscow no. 3U 25/4B (BM— holotype). [TLC: the 
lividic acid complex, atranorin.] 

Thallus corticolous, loosely attached, pale grey. Lobes rounded, up to 1 cm broad, 
abundantly ciliate, cilia 2-4 mm long. Upper side smooth, emaculate, more or less rugose 
towards the centre of the thallus. Medulla white. Underside black, with a broad, brown, 




Fig. 13 Parmelia indoafra Krog & Swinscow, T 6/1 1 1 (O). Rule = 1 cm. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



189 



naked marginal zone, rhizines sparse. Soralia and isidia absent. Apothecia submarginal, 
stipitate, thalline exciple and stipe strongly maculate, thalline margin dentate-ciliate, disc up 
to 1 cm in diameter, imperforate, spores 25-28 x 17-20//m, thick-walled. Conidia sub- 
lageniform, 7-9 jum long. 
TLC: the lividic acid complex, atranorin. 

The description is based on the Ugandan material. The specimens from West Africa were 
better developed, with broader lobes and longer cilia. 

Parmelia inexpectata is similar to P. amaniensis in most important morphological 
characters, and the two species differ mainly chemically. However, we consider the 
complexity of the lividic acid group of substances to warrant the acceptance of the chemical 
content as a species specific character in this case. For further discussion, see under 
'Circumscription of the species'. 

The species is not known outside Africa. In East Africa it is represented only by the type 
collection of P. paradoxa cited above. 




Fig. 14 Parmelia jacarandicola Krog & Swinscow, holotype (UPS). Rule = 1 cm. 



90 



H. KROG & T. D. V. SWINSCOW 



29. Parmelia jacarandicola Krog & Swinscow sp. nov. 



Fig. 14 



Thallus corticola, griseus. Lobi 1-2 cm lati, crenati, ciliis 0*5-2 (3) mm longis, superne leviter 
maculati. Soralia isidiaque nulla. Apothecia submarginalia, stipitata, disco perforate Spori 
15-16 x 8-10 /zm. Conidia filiformia, 10-15 /im longa. Acidum gyrophoricum, acidum psoromicum, 
et atranorinum continens. 

Thallus corticolous, loosely attached, pale grey. Lobes 1-2 cm broad, crenate, ciliate, cilia 
05-2 (3) mm long. Upper side smooth to pitted, rugose and cracked towards the centre of the 
thallus, faintly maculate. Medulla white. Underside black, with a broad, brown to mottled, 
naked marginal zone, rhizines fairly coarse, in scattered groups. Soralia and isidia absent. 
Apothecia submarginal, stipitate, up to 12 mm in diameter, thalline exciple rugose, exciple 
and stipe strongly maculate, thalline margin smooth, disc perforate, spores 1 5-16 x 8-10 jum, 
thin-walled. Conida filiform, 10-15 //m long. 

TLC: gyrophoric acid, psoromic acid (+), atranorin. 

Type: Kenya. Rift Valley, Nakuru [0° 17' S, 36° 5' E], on trunk of Jacaranda, 21 March 
1968, leg. Gunnar Gilenstam no. 2004 (UPS — holotype, O — isotype). 

Parmelia jacarandicola may superficially resemble P. maclayana, which, however, has 
sublageniform conidia and different chemical properties. Parmelia euneta differs in its larger 
spores, distinct maculae, and absence of psoromic acid. 

The production of psoromic acid is associated with the apothecia. Fertile lobes and the 
thalline exciple give a PD+ sulphur yellow reaction in the cortex, whereas the medulla reacts 
PD— . If a sterile lobe is tested with TLC, the presence of psoromic acid may go undetected. 

Parmelia jacarandicola may possibly be considered the parent morph of P. direagens, a 
sorediate species in which psoromic acid is associated with the soralia. However, there is a 
discrepancy in the conidia, which are sublageniform to rodshaped in P. direagens. An 
alectoronic acid producing strain which occurs in P. direagens is not yet known in P. 
jacarandicola. 




Fig. 15 Parmelia kwalensis Krog & Swinscow, holotype (O). Rule = 1 cm. 



P ARM ELI A (AMPHIGYMNIA) IN EAST AFRICA 



191 



East African records 

Kenya. Rift Valley Province, Nakuru District, Nakuru, Agricultural Show Ground, c. 1900 m, on a 
cultivated Jacaranda mimo si folia, Santesson 21663 b (UPS); Nakuru, on trunk of Jacaranda, 
Gilenstam 2004 (UPS, O— type collection). 

30. Parmelia kwalensis Krog & Swinscow sp. nov. Fig. 15 

Thallus cofticola, adnatus, virido-griseus. Lobi ad 1 cm lati, orbiculares, profunde divisi, crenati, 
ciliis 0*5-2 mm longis, superne maculati. Isidia ad laminam, ad centrum thalli aggregata. Apothecia 
et pycnidia ignota. Acidum norsticticum, acidum sticticum, acidum consticticum, et atranorinum 
continens. 

Thallus corticolous, adnate, green-grey. Lobes up to 1 cm broad, rounded, deeply divided, 
crenate, ciliate, cilia 0*5-2 mm long. Upper side distinctly maculate. Medulla white. 
Underside dark brown, with a narrow, pale brown, naked marginal zone, rhizines relatively 
long, slender, scattered almost to the margins. Soralia absent. Isidia laminal, crowded 
towards the centre of the thallus, cylindrical, simple or coralloid, some bearing cilia. 
Apothecia and pycnidia unknown. 

TLC: norstictic acid, stictic acid, constictic acid, atranorin. 

Type: Kenya. Coast Province, Kwale District, 2 km N of Gazi, 4° 22' S, 39° 30' E, in 
mangrove, February 1972, coll. H. Krog & T. D. V. Swinscow no. K 44/103 (O — holotype, 
BM — isotype). 

Parmelia kwalensis is considered to be the isidiate counterpart of the primary species P. 
aldabrensis, which grows in the same locality. 

The species recalls P. crinita in being isidiate and ciliate and producing stictic acid, but 
differs in the strongly maculate upper cortex, brown underside, and presence of norstictic 
acid. Furthermore, the two species have different ecological requirements, since P. crinita is 
a species of montane forests and inselbergs. Parmelia subtinctoria, which has a maculate 
upper cortex and a brown underside, differs both in chemical properties and ecology; it has 
not been collected below 1 500 m altitude in our area. 

Parmelia kwalensis is known only from the type locality where it grew on twigs in a 
mangrove by the Indian Ocean; we have collected it there on two different occasions. 

East African records 

Kenya. Coast Province, Kwale District, 2 km N of Gazi, K 44/103 (type collection), 3K 30/1 36. 

31. Parmelia leonis Krog & Swinscow sp. nov. Fig. 16 

Thallus corticola, griseus ad griseo-viridis. Lobi 0*5-1 (2) cm lati, orbiculares vel irregulariter incisi, 




Fig. 16 Parmelia leonis Krog & Swinscow, holotype (O). Rule = 1 cm 



192 H. KROG & T. D. V. SWINSCOW 

crenati, ciliis 0*3-2 mm longis, superne valde maculati, inferne plerumque omnino albi vel raro ad 
centrum nigri. Soralia isidiaque nulla. Apothecia marginalia vel submarginalia, stipitata, disco vulgo 
late perforate Spori 16-18 (20) x 8-10 jum. Conidia filiformia, 12-16(1 8) //m longa. Acidum proto- 
lichesterinicum et atranorinum continens. 

Thallus corticolous, loosely attached, pale grey to grey-green. Lobes 05—1 (2) cm broad, 
rounded or irregularly incised, crenate, ciliate, cilia 0*3-2 mm long. Upper side strongly 
maculate, cracked and rugose towards the centre. Medulla white. Underside entirely white, 
or rarely blackened in the centre with a wide white marginal zone, occasionally with brown 
blotches in unattached parts, strongly rugose, with white, pale brown, black-tipped, or black 
rhizines. Soralia and isidia absent. Apothecia marginal and submarginal, stipitate, up to 
15 cm in diameter, thalline exciple rugose, thalline margin entire or rarely dentate-ciliate, 
sometimes involute, disc mostly widely perforate, spores 16-18 (20) x 8-10 //m. Conidia 
filiform, 12-16(18) pm long. 

TLC: protolichesterinic acid, atranorin. 

Type: Kenya. Eastern Province, Machakos District, lava flow 5 km NW of Kibwezi, 
2°35 , S, 37° 5 TE, on shrubs, 1000 m, February \914,co\\.H.Krog& T.D. V. Swinscow no. 
3K 23/127 (O— holotype, BM, UPS— isotypes). 

Parmelia leonis differs from the protolichesterinic acid strain of P. abessinica in the almost 
entirely white underside and the longly filiform conidia. From P. glaucocarpoides Zahlbr. it 
differs in the same characters, and also in spores size which in P. glaucocarpoides is 
25-27 x 12-15 pm. Parmelia melanothrix (Mont.) Vainio, another primary species with 
protolichesterinic acid, has imperforate discs and spores 20-26 jum long, fide Hale (1965). 
Parmelia leonis often grows together with P. uberrima, which also has a white underside but 
has imperforate apothecia and contains alectoronic acid, and P. andina, which lacks cilia 
and reacts C+ blood red in the medulla (lecanoric acid). 

Parmelia leonis grows on twigs and thinner branches of shrubs in dry, well lit sites. It is 
locally abundant in the type locality at 1000 m and has further been collected once at 
1400 m in Kenya and once at 500-700 m in Tanzania. 

East African records 

Kenya. Eastern Province, Machakos District, 5 km NW of Kibwezi, 3K 23/127 (type collection), 
K20/25B, 120, 2K22/107, 5K2/30, Ol Doinyo Sapuk, 20 km SE of Thika, 2K 1/105. Tanzania. 
Eastern Province, Morogoro District, Kitulangalo Forest Reserve, 6 May 1978, Dahl(0). 

32. Parmelia lobulascens Steiner 

Verh. zool.-bot. Ges. Wien 53:234 (1903). — Parmotrema lobulascens (Steiner) Hale, Phytologia 

28 : 337 (1974). Type: Cameroons, Bornmiiller (WU — not seen). [TLC (Hale 1973): alectoronic acid, 

tf-collatolic acid, gyrophoric acid, atranorin.] 
Parmelia natalensis Steiner & Zahlbr., in Zahlbruckner, Bot. Jb. 60:515 (1926). — Parmotrema 

natalense (Steiner & Zahlbr.) Hale, Phytologia 28 : 337 (1974). Type: Natal, Drakens-Berge, Van 

Reenens-Pass, Schwarzer Berg, bei 1 700 m, Brunnthaler (W — not seen). 
Parmelia pseudonilgherrensis Asah., J. Jap. Bot. 29 : 370 (1954). — Parmotrema pseudonilgherrense 

(Asah.) Hale, Mycotaxon 5 : 441 (1977). Type: Korea, Mt Hokusui-Hakusan, Kankyo-Nando, 13 

August 1936, coll. U. Tsutani (TNS — lectotype). [TLC: alectoronic acid, a-collatolic acid, 

atranorin.] 

Thallus corticolous or saxicolous, often coriaceous, loosely attached, pale grey to grey-green, 
lobe margins sometimes tinged with brown. Lobes 0'5-L5(2)cm broad, rounded and 
crenate or irregularly incised-dentate, margins ascending or revolute, ciliate, cilia 
02-3 (5) mm long, coarse or slender, sometimes absent or poorly developed on peripheral 
lobes but present on lateral and central lobes, rarely entirely lacking. Secondary lobules 
common along the lobe margins and on the lamina. Upper side distinctly maculate, 
irregularly cracked, here and there with black, discoloured patches. Medulla white. 
Underside black, with a brown, naked marginal zone, rhizines fairly long, unevenly 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 193 

distributed. Soralia linear on ascending lobe margins or submarginal, pustular, and 
punctiform to confluent on revolute lobes. Apothecia uncommon, submarginal on 
peripheral or central lobes, usually distinctly stipitate on swollen stipes but sometimes sub- 
stipitate with a constricted base, thalline exciple strongly rugose, becoming sorediate, 
thalline margin crenate to lobulate, disc perforate or imperforate, often radially split, 
spores (13) 20-27 (30) x (8) 10-15 (18) /an, relatively thick-walled. Conidia filiform, 
(10) 12-16 (18) /zm long. 

TLC: alectoronic acid, ± or-collatolic acid, ± gyrophoric acid, atranorin. 

Hale (1965) found P. lobulascens morphologically very close to P. pseudonilgherrensis, but 
at the time the species was believed to contain gyrophoric acid only, while P. pseudo- 
nilgherrensis had alectoronic and a-collatolic acids. With the application of TLC, Hale 
(1973) found that P. lobulascens had both alectoronic and #-collatolic acids in addition to 
gyrophoric acid. Winnem (1975) gave the first report of apothecia in P. lobulascens, and on 
the basis of two fertile specimens she found that it had small spores, 13-17 x 8-10 /an. 
However, as in several secondary species, the apothecia of P. lobulascens produced few 
spores which often gave the impression of being deformed or abnormal (pyriform shape, 
extremely thin spore wall, shrunken cell content). The same phenomenon was encountered 
also in specimens of P. pseudonilgherrensis. We believe that the non-sorediate counterpart 
of P. lobulascens s. str., classified as P. diversa by Hale (1973) and Winnem (1975), gives 
better information on its normal spore size, namely 20-25 x 10-15 //m, which is within the 
variation range for P. pseudonilgherrensis. Since there is no important morphological, 
anatomical, or ecological differences between P. lobulascens and P. pseudonilgherrensis, we 
regard the gyrophoric acid as an accessory substance and reduce P. pseudonilgherrensis to 
synonymy with P. lobulascens. 

Parmelia natalensis has been regarded as distinct from P. lobulascens (as P. pseudo- 
nilgherrensis) mainly owing to its submarginal soralia on revolute lobes and its substipitate, 
imperforate apothecia centrally situated on the thallus, contrasting with the linear soralia on 
ascending lobe margins and the stipitate, perforate, submarginal apothecia of P. lobulascens. 
Unfortunately, the type of P. natalensis has not been available for study, but the exsiccate 
number 60 in Almborn, Lichenes Africani (LD), collected in Kenya, showed all the 
diagnostic characters mentioned in the protologue, including rather small spores. The 
specimen at first sight looked convincingly different from P. lobulascens, but when plentiful 
material was studied, no clear-cut discontinuities could be found. The centrally located 
apothecia of P. natalensis were not equivalent to laminal apothecia; they were merely 
submarginally situated on central lobes. In P. lobulascens the apothecia varied in individual 
specimens from perforate to entirely imperforate and radially split, while in P. natalensis the 
occasional perforate apothecium occurred in addition to the more common imperforate 
ones. Most specimens of the natalensis morphotype had spores in the interval 20-25 //m, an 
indication that the smaller spores sometimes seen (and cited, for example, in the protologue) 
were immature or abnormal. Specimens with abundant submarginal soralia on revolute 
lobes often had marginal soralia on ascending lobe margins on the same thallus. Some 
specimens with exclusively submarginal soralia even produced gyrophoric acid as an 
accessory substance and would have been included in P. lobulascens s. str. on chemical 
criteria. 

If we turn to the primary species, no parent morph corresponding to P. natalensis has been 
suggested in the literature, and it would indeed be extremely difficult to keep such a species 
separate from P. nilgherrensis. Within P. nilgherrensis it is possible to find specimens with 
substipitate rather than stipitate apothecia, or with apothecia mainly situated on central 
lobes, but none of these traits is suitable for the separation of species. We therefore regard P. 
natalensis as conspecific with P. lobulascens. 

Parmelia lobulascens is a common and widespread species in all four countries of our 
study, where it grows on both trees and rock throughout the montane forests, extending into 
the alpine zone; it has been collected between 1 800 and 3600 m altitude but is most common 



194 H. KROG & T. D. V. SWINSCOW 

above 2800 m. Outside our area it is known from South and West Africa, Nepal, and Korea 
(Hale, 1965), as well as from India (BM). 

Selected East African records 

Ethiopia. See Winnem (1975) under P. lobulascens, P. natalensis, and P. pseudonilgherrensis. Kenya. 
Central Province, Nyeri District, Mt Kenya, W slope, Naro Moru track (numerous coll. between 2200 
and 3500 m altitude), Aberdare Mountains, E of Lesatima, K25/8a. Rift Valley Province, Elgeyo 
Marakwet District, Cherangani Hills, 10-20 km S of Labot, 2K 7/14a, 2K 8/118, Sogotio Forest 8 km 
N of Chebiemit, 2K 10/106; Uasin Gishu District, 5 km NW of Timboroa Summit, 2K 19/1 18; 
Kericho District, Cheboswa ENE of Kericho 4K 17/117; Nakuru District, 1 km S of Mau Narok, 
4K 29/1 10. Eastern Province, Meru District, Mt Kenya, E side, at Themwe, 3K 16/164, Mt Kenya, N 
side, Sirimon track, 4K 20/124. Tanzania. Northern Province, Masai District, Ngorongoro Crater, 
Sipman 6371 A (herb. Sipman); Arusha District, Mt Meru Crater, T 5/126, Arusha National Park, near 
Ngurdoto gate, T 6/101. Uganda. Kigezi District, Bufumbira County, Muhavura, N side, U 16/10. 
Toro District, Busongora County, 10 km NW of Kilembe, slopes of Ruwenzori, 2U 13/2. Bugisu 
District, N Bugisu County, 05 km NW of ford over Sasa River, 2U 45/9, 18; Manjiya County, near 
Bukalesi, Lye L 13 (herb. Lye). 

33. Parmelia lophogena des Abb. 

Bull. Inst. fr. Afr. noire A, 20 : 19 (1958). — Parmotrema lophogenum (des Abb.) Hale, Phytologia 
28:337 (1974). Type: Guinee Franchise: Fouta-Djalon, a Dalaba, 1200 m, des Abbayes (US — 
isotype). [TLC: gyrophoric acid, fatty acids in the reddenda complex, atranorin.] 

Thallus corticolous, membranaceous, loosely attached, pale grey to green-grey. Lobes 
0*5-2 cm broad, sparsely crenate or crenate-lobulate, ciliate, cilia 2-^ (5) mm long. Upper 
side emaculate or faintly maculate, cortex fragile, cracking and flaking, pitted or rugose 
towards the centre. Medulla white. Underside black, with a wide, brown, naked marginal 
zone, rhizines sparse, in scattered groups. Coarse isidia and/or granular soredia present along 
the margins or in a broad marginal zone, laminal cilia not uncommon. Apothecia unknown. 
Conidia filiform, 12-15(17) //m long. 
TLC: gyrophoric acid, fatty acids in the reddenda complex, atranorin. 

We have previously discussed the diversity of the vegetative propagules in P. lophogena 




Parmelia louisianae Hale, K 20/1 19 (O). Rule 



P ARM ELI A (AMPHIGYMNIA) IN EAST AFRICA 195 

(Krog & Swinscow, 1975), and included P. diacidula Hale and P. dolosa des Abb. in its 
variation range. 

The species is known only from the African continent and Madagascar (Krog & Swinscow, 
1975, Fig. 1). In East Africa it has been collected in Kenya and Tanzania, where it grew in 
the lower montane forest between 1400 and 2400 m altitude. 

East African records 

Kenya. Rift Valley Province; Kericho District, Cheboswa ENE of Kericho, 4K 17/115; Laikipia 
District, Burguret, 4K 25/123; Narok District, Enabilibil, 22 km S of Siape bridge, 4K 31/108. Central 
Province, Nyeri District, Burguret River, 4K 27/109. Eastern Province, Marsabit District, Mt 
Marsabit, vicinity of Lake Paradise, 4K 6/138. 
For further records from Kenya and Tanzania, see Krog & Swinscow (1975). 

34. Parmelia louisianae Hale Fig. 17 

Phytologia 22 : 92 (\91\).—Parmotrema louisianae (Hale) Hale, Phytologia 28 : 337 (1974). Type: 
Louisiana, Desoto County, 3 mi N Sabine Co. Line on hwy 175, leg. M. E. Hale 34013 (US — 
holotype). [TLC: alectoronic acid, atranorin.] 

Thallus corticolous, loosely attached, pale grey to grey-green. Lobes 05—1 cm broad, 
rounded, margins ascending, crenate, ciliate, cilia 0*5-2 mm long. Upper side maculate, 
rugose towards the centre. Medulla white. Underside black in the centre, with a broad, white 
marginal zone, lobe-ends sometimes with brown blotches. Soralia submarginal on revolute 
lobes. Apothecia not seen. Conidia shortly filiform, 10-12 //m long. 
TLC: alectoronic acid, ± #-collatolic acid, atranorin. 

This rare lichen has previously been known from two localities in Louisiana and one in 
Virginia (W. Culberson, 1973). We have made four small collections of it, all from the same 
locality in Kenya. Both Hale {\91\b) and W. Culberson (1973) considered that P. louisianae 
was the sorediate counterpart of the American species P. rigida Lynge. Our specimens grew 
together with P. uberrima and might be interpreted as the sorediate counterpart of that 
species. The two primary species have similar chemical properties except for the addition of 
norstictic acid in some specimens of P. rigida, and they have both got a mainly white 
underside. However, P. rigida differs from P. uberrima in having perforate apothecia and 
longly filiform conidia. The sorediate species has never been found fruiting. No conidia were 
found in the two American specimens at our disposal, but the Kenyan specimens had the 
same shortly filiform conidia as P. uberrima. As long as P. louisianae is known only in a 
sterile state, its parent morph remains uncertain, and it is not conclusively settled that the 
American and African populations belong to the same species. 
All our specimens grew on shrubs in a thicket in a dry, well lit site at 1000 m altitude. 

East African records 

Kenya. Eastern Province, Machakos District, lava flow 5 km NW of Kibwezi, K20/1 19, 2K 22/109, 

3K23/122, 123. 



35. Parmelia maclayana Miill. Arg. Fig. 18 

Flora, Jena 14:316 (1891). — Parmotrema maclayanum (Miill. Arg.) Hale, Phytologia 28:337 
(1974). Type: Hb. Maclay, ex Afric. capric. [? collector] (G — holotype, BM — isotype). [TLC: 
alectoronic acid, unknown with alectoronic acid, atranorin.] 

Parmelia procera Steiner & Zahlbr., in Zahlbruckner, Bot. Jb. 60:537 (1926). — Parmotrema 
procerum (Steiner & Zahlbr.) Hale, Phytologia 28 : 338 (1974). Type: [Tanzania] E Usambara, near 
Amani, Brunnthaler (US — isolectotype). [TLC: alectoronic acid, <*-collatolic acid, atranorin.] 

Parmelia breviciliata Hale, Contr. U.S. natn. Herb. 36 (5) : 282 (1965). — Parmotrema breviciliatum 
(Hale) Hale, Phytologia 28 : 335 (1974). Type: Transvaal, Distr. Pilgrims Rest, 4 miles SE of P. R., 
on rocks near road, leg. Ove Almborn 7800 (LD — holotype). [TLC: alectoronic acid, unknown with 
alectoronic acid, atranorin. 1 



96 



H. KROG & T. D. V. SWINSCOW 




Fig. 18 Parmelia maclayana Mull. Arg., Nordal INB 849 (O). Rule = 1 cm. 



Thallus corticolous or saxicolous, coriaceous, pale grey. Lobes (0 - 5) 1-2 (2*5) cm broad, 
entire or crenate-dentate, ciliate, cilia sparse, 0'5-2 (3) mm long. Upper side emaculate, 
irregularly cracked towards the centre. Medulla white. Underside smooth or rugose, black in 
the centre, with a dark brown marginal zone, rhizines of medium length, in scattered groups, 
sometimes coming close to the margins. Soralia and isidia absent. Apothecia submarginal, 
stipitate, up to 3 cm in diameter, thalline exciple more or less rugose, maculate, thalline 
margin thin, smooth or weakly crenate, disc imperforate or with a small perforation, spores 
12-1 5x7-10 p.m. Conidia sublageniform, 6-7 jum long. 

TLC: (1) alectoronic acid, unknown with alectoronic acid, atranorin; (2) alectoronic acid, 
tf-collatolic acid, atranorin. 

Parmelia maclayana has been separated from P. procera mainly by its perforate apothecia. 
However, this is a variable feature in P. maclayana; the only apothecium in one of the type 
specimens, for example, is imperforate. There is a small chemical difference between them, 
but it is too slight to warrant the acceptance of two different species. Parmelia breviciliata 
differs from P. maclayana only in being saxicolous. The broad lobes, huge apothecia, small, 
thin-walled spores, and sublageniform conidia are the same in all three taxa. Chemotype (2) 
is represented only by the type of P. procera and by the specimen Greenway 1001 cited 
below; the rest of the specimens belong in chemotype ( 1 ). 

In East Africa Parmelia maclayana has been collected in miombo woodland, artificial 
habitats, and well lit sites in the lower montane forest between c. 900 and 2200 m altitude. 
We have also seen specimens from Sierra Leone, Cameroon, Zambia, Zimbabwe and 
Malawi. Hale (1965, under P. breviciliata, P. procera, and P. maclayana) cited numerous 
collections from continental Africa as well as a few from Madagascar, Thailand, Brazil, and 
New Caledonia. 

East African records 

Kenya. Eastern Province, Aberdare Mountains, Kimakia Forest Station, Ryvarden 9032 (O). 
Tanzania. E. Usambara, Amani, Greenway 1001 (BM); E. Usambara, near Amani, Brunnthaler (type of 
P. procera). Lake Province, Bukoba, 11/92, Stuhlmann 4059 (BM, sub Parmelia nitens Mull. Arg.). 
Southern Province, Songea District, Gumbiro, in miombo woodland, Nordal INB 849 (O); Songea 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 197 

District, Kigonsera, Missionsstation bei Peramiko, Dietrich (M). Western Province, Kigoma District, 
Kungwe-Maheli Peninsula, Kungwe Mountains, September 1959, Harley (BM, sub P. aldabrensis). 
Uganda. West Mengo District, Busiro County, Entebbe, on epiphytic orchids, Proctor s.n. (BM); 
Naiguru, Dale L 49 p. p. (BM). 

36. Parmelia mellissii Dodge 

Ann. Mo. bot. Gdn 46: 134 (1959).— Parmotrema mellissii (Dodge) Hale, Phytologia 28 : 337 
(1974). Type: St Helena, coll. Melliss 23 (BM— holotype). [TLC (Winnem 1974): alectoronic acid, a- 
collatolic acid, atranorin.] 

Thallus corticolous, membranaceous or rarely somewhat coriaceous, loosely attached, pale 
grey to grey-green. Lobes 0-3-1 cm broad, margins ascending, crenate, abundantly ciliate, 
cilia slender, 2-\ mm long. Upper side emaculate, pitted or weakly rugose, cortex fragile, 
cracking and often flaking, especially near the lobe margins. Medulla white, with patches of 
an ochraceous, K+ purple pigment (skyrin). Underside black, with a narrow, dark brown 
marginal zone, rhizines long, slender, in scattered groups. Soralia marginal and submarginal, 
here and there interspersed with coarse, ciliate isidioid outgrowths which become sorediate 
with time, soredia granular. Mature apothecia and pycnidia not seen in East African 
material. (According to Hale (1965) the apothecia are imperforate, with spores 16-22 x 10- 
14 jum.) 
TLC: alectoronic acid, #-collatolic acid, ± skyrin, atranorin. 

Parmelia mellissii recalls P. cryptoxantha and P. lophogena in its fragile and flaking upper 
cortex and long cilia. However, P. cryptoxantha has mainly laminal pustules and open 
dactyls with sparse production of soredia, and its medulla is pigmented. Parmelia lophogena 
is a larger, more broad lobed species which differs mainly chemically; it seems to be 
restricted to Africa. 

Parmelia mellissii grows in the lower montane forest, where it has been collected between 
2000 and 2400 m altitude. The species is widespread in the Americas and is known from 
India, the Canary Islands, and St Helena (Hale, 1965), as well as from New Zealand (BM). In 
East Africa it is rare. 

East African records 

Kenya. Central Province, Nyeri District, Mt Kenya, above entrance to National Park, 2K 32/4. Eastern 
Province, Meru District, Mt Kenya, E side, at Themwe, 3K 16/129. Tanzania. Northern Province, 
Arusha District, Mt Meru, confluence of River Jekukumia with River Ngare Nanyuki, T 11/5, 6; 
Moshi District, Kilimanjaro, Mweka, Sitari 280 (TUR). 

37. Parmelia nilgherrensis Nyl. 

Flora, Jena 52:291 (1869).— Parmotrema nilgherrense (Nyl.) Hale, Phytologia 28:338 (1974). 
Type: Ind. Or., in montib. nilgherrensibus, leg. Perrottet (H-NYL 35337 — holotype). [TLC: 
alectoronic acid, #-collatolic acid, atranorin.] 
Parmelia diversa Hale, Phytologia 27: 1 (1973). — Parmotrema diversum (Hale) Hale, Phytologia 
28:336 (1974). Type: Ethiopia, Bale Prov., mountain pass between Adaba and Goba, 3600 m, 
January 1972, coll. H. Krog E 22/19 (US— holotype, O— isotype). [TLC: alectoronic acid, 
tf-collatolic acid, gyrophoric acid, atranorin.] 

Thallus corticolous or saxicolous, usually coriaceous, loosely attached, pale grey, often with 
a brownish tinge towards the margins. Lobes 1-1 '5 (2) cm broad, rounded and crenate or 
irregularly incised-dentate, margins often ascending, ciliate, cilia 02-3 mm long, slender to 
coarse. Upper side strongly maculate, cracked, sometimes with blackened patches. Medulla 
white. Underside black, with a brown, naked marginal zone, rhizines slender to coarse, of 
varying length and density, often penicillate. Soralia and isidia absent. Apothecia up to 
2*5 cm in diameter, submarginal, stipitate on often swollen stipes, more rarely substipitate 
with a constricted base, thalline exciple strongly rugose, thalline margin dentate, involute, 
disc perforate or imperforate, spores (20) 23-27 (30) x 10-15 //m, fairly thick-walled. 
Conidia filiform, (10) 12-14 (16) /am long. 



198 



H. KROG & T. D. V. SWINSCOW 



TLC: alectoronic acid, ± or-collatolic acid, ± gyrophoric acid, atranorin. 

Saxicolous forms at high altitudes are extremely coriaceous, with short, coarse cilia, while 
corticolous specimens in the montane forests usually have a thinner thallus and longer, more 
slender cilia. This variation seems to be environmentally induced and is not correlated with 
the presence or absence of gyrophoric acid. Laminal lobules, which were one of the 
characteristic features of P. lobulascens Steiner, the putative sorediate morph of P. diversa, 
are found in specimens with or without gyrophoric acid. The intermediate spores with a 
rather thick spore wall as well as the filiform conidia are the same for both chemotypes. We 
consider gyrophoric acid to be an accessory substance in this case and regard P. diversa as 
conspecific with P. nilgherrensis. 

Parmelia maclayana differs from P. nilgherrensis in its emaculate upper cortex, smaller 
spores, sublageniform conidia, and different ecological requirements. 

Parmelia nilgherrensis is a species of the montane forests, the ericaceous zone, and the low 
alpine zone; it has been collected between 2000 and 3600 m altitude, but is most common 
above 2800 m. The gyrophoric acid strain is most widespread in Ethiopia. We have seen 
only two additional specimens, one from Kenya and one from Tanzania. Outside our area 
the species is known from Sikkim, India, Sri Lanka, China, and Thailand (Hale, 1965, Fig. 
28). 

Selected East African records 

Ethiopia. Bale Province, Gasore Summit, Tapper 960a, 962 (BM). Begemder Province, Semen Nat. 
Park, Geech plateau, Heinonen 1 187 (H). Chokke Mts, north of Debra Marcos, C. B. E. E. 34 (BM). 
For further Ethiopian records, see Winnem (1975) under P. diversa and P. nilgherrensis. Kenya. Central 
Province, Nyeri District, Mt Kenya, W side, Naro Moru track, K 28/122, 520, Aberdare Mountains, 
near Karuru-Gura waterfalls, 3K 32/1 14; Kirinyaga District, Mt Kenya, S side, 2 km NW of Irangi 
Forest Station, K 48/1 38. Eastern Province, Meru District, Mt Kenya, E side, at Themwe, 3K 16/167; 
Rift Valley Province, Elgeyo Marakwet District, Cherangani Hills 10 km S of Labot, 2K 8/190; 
Trans-Nzoia District, Mt Elgon crater at Maji ya Moto (hot spring), Hedberg 912 (UPS). Tanzania. 
Northern Province, Arusha District, Mt Meru Crater, T 5/143. Uganda. Bugisu District, N Bugisu 
County, by Sasa Hut on Mt Elgon, 2U 44/8. Toro District, Busongora County, Mt Ruwenzori, near 
Nyamileju Hut, Lye L 6 1 (herb. Lye). 

38. Parmelia paraphypoptropa W. Culb. Fig. 19 

Bryologist 76:29 (1973). — Parmotrema paraphypotropum (W. Culb.) Hale, Phytologia 28:338 




Fig. 19 Parmelia parahypotropaW . Culb., 3 K 30/107(0). Rule= 1 cm. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



199 



(1974). Type: Formosa, Kural, 250 m, Kurokawa 1487 (TNS — holotype). [TLC: norstictic acid, 
galbinic acid, salazinic acid, atranorin.] 

Thallus corticolous, loosely attached, pale green-grey. Lobes up to 06 cm broad, deeply 
divided, laciniae 1*2-1 '5 mm broad, sublinear, dichotomously branched, ciliate, cilia 
1-2 mm long. Upper side faintly to distinctly maculate, often cracked. Medulla white. 
Underside black in the centre, with a white or mottled marginal zone mainly restricted to the 
laciniae, rhizines short, sparse, in the centre of the thallus. Soralia labriform at the tips of the 
laciniae, soredia farinose. Apothecia and pycnidia not seen. 
TLC: norstictic acid, galbinic acid, salazinic acid, ± fatty acid (trace), atranorin. 

The narrow laciniae and labriform soralia distinguish P. parahypotropa from all other 
Amphygmnia species in our area. 

In East Africa P. parahypotropa is restricted to the coast of the Indian Ocean, where it has 
been collected on twigs in mangroves and on the trunk of a Cocos at 300 m altitude. It has 
been reported from similar habitats in Australia by Stevens & Rogers (1979). 

East African records 

Kenya. Coast Province, Kwale District, Kwale, Forestry Station, Santesson 20802b (UPS), 2 km N of 

Gazi, K 44/34, 1 10, 3K 30/107; Kilifi District, Mida Creek, 3K 29/1 15. 



39. Parmelia pardii Krog & Swinscow sp. nov. 



Fig. 20 



Thallus saxicola, griseus. Lobi 0-8-1-5 (2) cm lati, orbiculares, integri vel leniter crenati, eciliati, 
superne nitidi, emaculati. Soralia ad lobos laterales, marginalia et linearia, ad laminam punctiformia 
vel subcapitata, vulgo confluentia. Apothecia laminalia, disco primum imperforato demum 
perforate Spori 10-13 x 6-8 jum. Conidia filiformia, 10-14 //m longa. Acidum fumarprotocetra- 
ricum, acidum protocetraricum, et atranorinum continens. 

Thallus saxicolous, adnate, pale grey. Lobes 0-8-1-5 (2) cm broad, rounded, entire or 




Fig. 20 Parmelia pardii Krog & Swinscow, holotype (O). Rule = 1 cm. 



200 H. KROG & T. D. V. SWINSCOW 

weakly crenate, eciliate. Upper side shiny, emaculate, more or less cracked and rugose 
towards the centre. Medulla white. Underside black, with a broad, brown, naked marginal 
zone, rhizines fairly coarse, mainly black but occasionally brown peripherally, peripheral 
papillae sometimes present. Soralia marginal and linear on lateral lobes and punctiform to 
subcapitate on the lamina, often confluent over larger areas, soredia granular. Apothecia 
laminal, thalline exciple sorediate, disc at first imperforate, becoming perforate with age, 
spores 10-13 x 6-8 /im. Conidia filiform, 10-14 jum long. 

TLC: fumarprotocetraric acid, protocetraric acid, atranorin. 

Type: Kenya. Eastern Province, Machakos District, Lukenia, 30 km SE of Nairobi, 
1° 28' S, 37° 04' E, on rock and over plant remains, 1750 m, February 1974, coll. H. Krog & 
T. D. V. Swinscow no. 3K 1/102 (O— holotype; BM, UPS— isotypes). 

Parmelia pardii recalls P. pseudograyana in chemical properties and in its saxicolous habit. 
However, that species has marginal cilia, imperforate apothecia, and sublageniform conidia. 
Morphologically P. pardii is close to P. defecta, with which it sometimes grows. The 
apothecia of P. defecta are not known, but in its supposed parent morph P. soyauxii they are 
distinctly perforate. That, and the considerable difference in chemistry, made us decide not 
to include P. pardii in P. defecta as a chemical strain. Fumarprotocetraric acid is the main 
diagnostic compound of P. pardii, while the much more commonly occurring protocetraric 
acid is present in lower concentrations 

Parmelia pardii is a species of exposed acidic rock. It has been collected at 1650-1750 m 
altitude. 

East African records 

Kenya. Eastern Province, Machakos District, Lukenia, 30 km SE of Nairobi, 3K 1/102 (type 
collection). Coast Province, Taita District, N of Mwanda summit, 2K 28/107; Taita District, E of 
Mwanda summit, 2K 29/102. 

40. Parmelia perlata (Huds.) Ach. 

Meth. Lick. : 2 16 (1803). — Lichen perlatus Huds., Fl. Angl. : 448 (1762). — Parmotrema perlatum 
(Huds.) Choisy, Bull. mens. Soc. linn. Lyon 21 : 174 (1952). Type: Specimen and pi. 20, fig. 39B, 
Dillenius, Hist. Muse. : 147 (1742 ['1741 ']) (OXF— lectotypotype, not seen). 

Thallus corticolous, membranaceous, loosely attached, pale grey to grey-green. Lobes 
0*5—1 -2 cm broad, margins ascending, crenate, ciliate, cilia 0*5-1 -5 mm long. Upper side 
emaculate to faintly maculate, pitted and uneven, often cracked. Medulla white. Underside 
black in the centre, with a brown marginal zone, densely rhizinate, rhizines short, sometimes 
extending almost to the margins. Soralia marginal, linear to subcapitate on somewhat 
revolute lobes, sometimes spreading submarginally. Apothecia and conidia not seen in East 
African material. (According to Hale (1965) the apothecia are imperforate with spores 
22-30 x 13-1 6 //m.) 

TLC: stictic acid, constictic acid, atranorin. 

Parmelia perlata is closely related to P. bangii, a species which differs in having 
submarginal, pustular soralia and a fragile and flaking upper cortex. 

In East Africa we have collected P. perlata in inselbergs, montane forests, and the 
ericaceous zone between 1400 and 3100m altitude. The species is pantemperate and 
widespread in both the northern and southern hemispheres (Hale, 1965, Fig. 20). 

East African records 

Ethiopia. Sidamo Province, between Kebre Mengist and Wadera, Winnem 595/5, 6 (O). Kenya. 
Central Province, Nyeri District, Mt Kenya, W side, above entrance to National Park, K 32/1 10, Mt 
Kenya, Naro Moru track, upper forest, 3100 m, K 35/105, Aberdare Mts, 10 km W of Tusha, 
3K 31/104. Rift Valley Province, Nakuru District, Londiani Forest SE of Mau Summit, 2K 5/106. 
Eastern Province, Marsabit District, vicinity of Lake Paradise, 4K 6/139. Tanzania. Northern 
Province, Arusha District, Mt Meru, confluence of River Jekukumia with River Ngare Nanyuki, 
T 11/114, 128. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 201 

41. Parmelia permutata Stirton 

Scott. Nat. 4 : 252 (\%ll-l%).—Parmotremapermutatum (Stirton) Hale, Phytologia 28 : 338 (1974). 
Type: Queensland, Brisbane, /. M. Bailey 25 (BM — holotype). [TLC: gyrophoric acid, pigment, 
atranorin.] 

Thallus corticolous, loosely attached, pale grey to grey-green. Lobes 0*5-1 *5 cm broad, 
weakly crenate, ciliate, cilia 1-3 (4) mm long. Upper side emaculate, smooth to faintly 
pitted, cracked in older parts. Medulla yellow, ochraceous, or salmon pink, pigment K— . 
Underside black, with a broad, brown or mottled, naked marginal zone, rhizines sparse, in 
groups. Soralia marginal, linear, often spreading submarginally. Apothecia absent. Pycnidia 
present but conidia not found. 
TLC: gyrophoric acid, pigment, atranorin. 

If the medullary pigment is overlooked, P. permutata may be confused with P. sancti- 
angelii, another emaculate, sorediate species with gyrophoric acid. However, in P. 
permutata the gyrophoric acid is concentrated near the upper cortex, giving a red reaction 
upon application of C, while the rest of the medulla remains yellowish pink. 

In East Africa P. permutata is a species of artificial habitats and well lit sites 
in the lower montane forest. It has been collected between 1100 and 2400 m altitude in 
all four countries of our study. Outside Africa it is widespread, being known from Australia 
(type), India, Sumatra, and Haiti (Hale, 1965), Papua New Guinea (Kurokawa, 1979), and 
Brazil (W, O). 

East African records 

Ethiopia. See Winnem (1975). Kenya. Eastern Province, Machakos District, Ol Doinyo Sapuk, 20 km 
SE of Thika, 2K 4/105. Coast Province, Taita District, W of Wundanyi, 2K 25/103. Rift Valley 
Province, Kericho District, Kericho Tea hotel, 4K 7/106. Tanzania. Northern Province, Arusha 
District, Arusha National Park, Juniper Hill, T 3/120. Uganda. East Mengo District, Kyagwe County, 
Owen Falls Dam, facing Jinja, U 4/15. Toro District, Burahya County, street in Fort Portal, 2U 17/11. 
Kigezi District, Kinkizi County, 30 miles NW of Kabale, Impenetrable Forest, Burnet AMB 2 1 3 (BM). 



42. Parmelia pigmentifera Krog & Swinscow sp. nov. Fig. 21 

Thallus corticola, griseus. Lobi 0* 5-0* 8 cm lati, ciliis 0*5-2(3) mm longis, superne emaculati. 
Medulla pigmento ochraceo, K— . Soralia isidiaque nulla. Apothecia marginalia vel submarginalia, 
stipitata, margine thallino laevi ad leniter dentato, ciliato, disco imperforato. Spori 18-22 (24) x 
10-12 (14) //m. Conidia leniter sublageniformia, 7-8 jum longa. Acida ignota, pigmentum, et 
atranorinum continens. 

Thallus corticolous, loosely attached, pale grey. Lobes 0*5-0*8 cm broad, ciliate, cilia 
0*5-2 (3) mm long. Upper side emaculate, often pruinose, smooth, becoming rugose with 
age. Medulla pigmented, pigment pink to ochraceous, K— . Underside black, with a brown, 
naked marginal zone, rhizines sparse. Soralia and isidia absent. Apothecia marginal and 
submarginal, stipitate, stipe often formed by a convoluted lobe, thalline exciple and stipe 
strongly maculate, thalline margin smooth to weakly dentate, ciliate, disc up to 1 cm in 
diameter, imperforate, spores 18-22 (24) x 10-12 (14) jum. Pycnidia numerous, conidia 
weakly sublageniform, 7-8 fim long. 

TLC: two undetermined substances in the following Rf classes. TDA: 2-3, 4; HEF: 4-5, 6; 
and TA: 2-3, 5-6, giving a strong bluish white fluorescence under long- wave UV light; 
undetermined pigment, atranorin. 

Type: Kenya. Coast Province, Kilifi District, Mida Creek, edge of mangrove, 3° 22' S, 
39° 57' E, February 1974, coll. H. Krog & T. D. V. Swinscow no. 3K 29/1 12 (O— holotype; 
BM,UPS— isotypes). 

Most primary Amphigymnia species with a pigmented medulla differ from P. pigmentifera, 
among other characters, in having larger spores. An exception is P. subcolorata (also 



202 



H. KROG & T. D. V. SWINSCOW 




Fig. 21 Parmelia pigmentifera Krog & Swinscow, holotype (O). Rule = 1 cm. 

occurring in East Africa), which, however, has perforate apothecia, filiform conidia, and 
different ecological requirements. 

Parmelia pigmentifera grows on twigs in mangroves on the Kenyan coast, and has also 
been collected at 70 m altitude near the coast in Tanzania. 

East African records 

Kenya. Coast Province, Kilifi District, Mida Creek, 3K 29/1 12 (type collection). Tanzania. Eastern 

Province, Rufiji District, Ngulakula, in miombo woodland, Nordal INB 758 (O). 



43. Parmelia pilosa Stizenb. 

Ber. Tat. St Gall, naturw. Ges. 1888-89 : 165 (\890).—Parmelina pilosa (Stizenb.) Hale, Phytologia 
28:483 (1974). Type: Republic Oranje, saxicola in montibus trachyticis Taaibosch-Kranz ad 
Rhenoster-Rivier, Rehmann (ZT — lectotype, not seen). 

Thallus corticolous or rarely saxicolous, usually adnate, pale grey to slate grey. Lobes deeply 
divided, 0"5-l(l*5)cm broad, crenate, ciliate, cilia coarse, 0-2-1 (2) mm long, often 
branched. Upper side distinctly maculate, sometimes partly pruinose. Medulla white. 
Underside black, with a brown marginal zone, usually rhizinate to the margins, rhizines 
dimorphous, in part long and coarse, in part short and slender. Soralia laminal to sub- 
marginal, punctiform. Apothecia not seen in East African material. Conidia filiform, 
12-16//m long. 
TLC: undetermined fatty acid, atranorin. 

Parmelia consors Nyl., not known from East Africa, is considered to be the parent morph of 
P. pilosa (Hale, 1976/?). Since it has partly perforate apothecia and filiform conidia we 
believe that the species pair is better placed in Amphigymnia than in genus Parmelina Hale 
(=subgenus Cyclocheila (Vainio) Krog & Swinscow). Species with dimorphous rhizines 
extending to the margins, such as P. subsumpta, are already accommodated in 
Amphigymnia. 

Parmelia pilosa has been collected on trunks and branches of solitary trees, often in 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 203 

artificial habitats, rarely on rock, in well lit sites between 1300 and 2000 m altitude. Outside 
our area it is known from South Africa and South America (Hale, \916b). 

East African records 

Ethiopia. Shewa Province, Debre Zeit, near Green Lake, Winnem 159/12 (O); Wondo Gennet, 20 km 
S of Shashemenne, E 5/30, Winnem 494/12 (O). Sidamo Province, E shore of Lake Awasa, E 6/8, 
Winnem 415/12, 442/10 (O); Awasa, NLM resort, Winnem 492/8 (O); Yirga Alem, Sidamo Provincial 
Hospital, E 20/21; Wadera, Winnem 510/18, 512/18, 596/17 (O). Kenya. Coast Province, Taita 
District, vicinity of Wundanyi 2K25/106, 2K26/103, 2K27/137. Rift Valley Province, Samburu 
District, WSW slope of Warges, 4K 3/115; Kericho District, Kericho Tea Hotel, 4K 16/105. Nyanza 
Province, Kisii District, 6 km E of Keroka, 4K 1 1/107, Kisii, 4K 13/104. Uganda. Mubende District, 
Buwekula County, Mubende Hill, 3U 12/3. Masaka District, Buddu County, near Kurumba, Lye 
L 222 B (herb. Lye). 

44. Parmelia planatilobata Hale 

/. Jap. Bot. 40 : 200 (1964).— Parmotrema planatilobatum (Hale) Hale, Phytologia 28 : 338 (1974). 
Type: Mountain Garden of Tjibodas, Java, c. 1400 m, Kurokawa 2 160 (TNS — holotype, not seen). 

Thallus corticolous or saxicolous, loosely attached, pale grey. Lobes 0*5-0*8 cm broad, 
rounded, entire or crenate, ciliate, cilia slender, 1-2 mm long. Upper side emaculate. 
Medulla white, with patches of an ochraceous, K+ purple pigment (skyrin). Underside black, 
with a brown, naked marginal zone, rhizines long, slender. Soralia absent. Dorsiventral, 
palmate, ciliate lobules and coralloid, more or less flattened, ciliate isidia abundant near the 
margins of the lobes, in part also marginally situated. One immature apothecium seen, no 
spores or conidia. 
TLC: gyrophoric acid, ± skyrin, atranorin. 

Parmelia planatilobata is apparently rare in East Africa. There are three collections of it, 
one from rock and two from tree bark. They all grew between 800 and 1000 m altitude in the 
Usambara Mountains. The species was previously known from Java and Malaya (Hale, 
1965). 

East African records 

Tanzania. Tanga Province, Lushoto District, Usambara Mts, Amani, Dodwe (road towards Derema), 
Santesson 23360 (UPS), Amani, in the surroundings of Forestry House, Vitikainen 9308 (H), Amani 
Forest Reserve, T. & S. Pocs 6 102/V-C (herb. Pocs). 



45. Parmelia poolii Dodge Fig. 22 

Ann. Mo. bot. Gdn 46 : 146 (1959). Based on Parmelia proboscidea var. sorediifera Mull. Arg., Flora, 
Jena 67:615 (1884), non Parmelia soredifera Sprengel (1827). Type: Central Madagascar, 
Hildebrandt s.n. (G — lectotype). [TLC: alectoronic acid, unknown with alectoronic acid, atranorin.] 

Thallus corticolous or saxicolous, loosely attached, pale grey. Lobes 0*5-2 cm broad, 
rounded, entire to crenate-dentate, ciliate, cilia sparse to abundant, 0*3-2 (4) mm. Upper 
side emaculate, shiny, finely cracked over most of the thallus. Medulla white. Underside 
black, with a dark brown, naked marginal zone, rhizines short to medium, in scattered 
groups. Soralia mainly lateral, marginal, linear to subcapitate. Apothecia not seen. Conidia 
sublageniform, 7-8 jum long. 

TLC: (1) alectoronic acid, unknown with alectoronic acid, atranorin; (2) alectoronic acid, 
tf-collatolic acid, atranorin. 

The type specimen of P. poolii was saxicolous and obviously represents the sorediate 
counterpart of P. breviciliata Hale, which we have reduced to synonymy with P. maclayana 
on the basis of spore and conidium characters (see above under P. maclayana). 

Parmelia poolii was considered conspecific with P. rampoddensis Nyl. by Hale (1965). 
However, type specimens of P. rampoddensis (H, S) indicate a fairly small species with 



204 



H. KROG & T. D. V. SWINSCOW 




Fig. 22 Parmelia poolii Dodge, 2K6/1 15 (O). Rule = 1 cm. 



marginal soralia on involute lobes, abundant laminal soralia, a pale, mottled marginal zone 
on the underside, and abundant cilia which produce a purple pigment never seen in East 
African material. No conidia were found in the type specimens, but a specimen from Sri 
Lanka, Pidurutalagala, 12 January 1972, Dahl (O), corresponding to the type in every 
respect, produced rod-shaped conidia c. 6 fim long. We doubt whether P. rampoddensis 
occurs in East Africa. The specimens cited from Ethiopia by Winnem (1975) are too scrappy 
for positive identification. 

Parmelia arnoldii Du Rietz is another emaculate, ciliate, sorediate Amphigymnia species 
with alectoronic and <ar-collatolic acids. It differs from our specimens mainly in a more 
membranaceous thallus, which cannot be considered an important character. However, 
those of our specimens which produced pycnidia had sublageniform conida 7-8 jum long, 
while European specimens of P. arnoldii produced filiform conidia 10-12 (14) jam long. We 
therefore include our specimens in P. poolii. Parmelia arnoldii does not seem to occur in 
East Africa. 

Parmelia lobulascens differs from P. poolii in its strongly maculate upper cortex, longly 
filiform conidia, and accessory gyrophoric acid. 

Parmelia poolii has been collected in the lower montane forest in Kenya between 2000 
and 2300 m altitude, and in one locality in Tanzania at 900 m. Outside our area we have 
seen specimens from Thailand (Kurok. Lich. Rar. Exs. no. 36, O), Madagascar (type 
collection), and Australia (O). Owing to taxonomic confusion its general distribution is not 
known, but it seems to agree well with that of P. maclayana. 

East African records 

Kenya. Rift Valley Province, Elgeyo Marakwet District, 7 km N of Chebiemit, 2K 6/1 15. Central 

Province, Kirinyaga District, Mt Kenya, S slope, 2 km NW of Irangi Forest Station near River Ena, 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 205 

K 48/124; Nyeri District, Mt Kenya, S slope, Regati Forest Station, Ryvarden 9822b, 9825 (O). Eastern 
Province, Meru District, Mt Kenya, E slope, on track to Themwe, 3K 15/1 16. Tanzania. Tanga 
Province, Lushoto District, Usambara Mountains, Amani, in the surroundings of Forestry House, 
Santesson 32153 (UPS). 

46. Parmelia praesorediosa Nyl. 

Sert. Lich. trop. Labuan Singapore: 18 (1891). — Parmotrema praesorediosum (Nyl.) Hale, 
Phytologia 28 : 338 (1974). Type: Singapore, 30 November 1879, E. Almquist (H-NYL 35547— 
holotype, S — isotype). [TLC: fatty acids in the reddenda group, atranorin.] 
Parmelia mordenii Hale, Smithson. Contr. Bot. 4 : 19 (1971). — Parmotrema mordenii (Hale) Hale, 
Phytologia 28:337 (1974). Type: Dominica, north of Coulibistri, c. 30 m, Hale 35649 (BM— 
isotype). [TLC: fatty acids in the reddenda group, atranorin.] 

Thallus corticolous or saxicolous, adnate, pale grey. Lobes 0*4-0*8 (1) cm broad, rounded, 
entire or crenate, eciliate, sorediate margins ascending. Upper side dull, emaculate, slightly 
rugose and cracked in older parts. Medulla white. Underside black, with a white, mottled, or 
brown marginal zone, rhizines short, sparse. Soralia marginal, linear to sublabriform, some- 
times spreading on to the lamina and then punctiform to subcapitate. Apothecia not seen in 
East African material. (According to Hale (1965) the apothecia are imperforate and the 
spores 1 5-2 1 x 7-10/zm.) Pycnidia rare, conidia sublageniform, 7-8 jum long. 
TLC: fatty acids in the reddenda group, atranorin. (Medulla K+ yellow.) 

Hale (1971a) distinguished P. praesorediosa from P. mordenii by its corticolous habit, 
generally smaller, shorter lobes, thinner, more greenish thallus, finer soredia often produced 
in linear, crescent-shaped, or labriform soralia without formation of subfatiscent structures, 
and the K— or faint K+ yellow reaction in the medulla (against K+ yellow in P. mordenii). 
We have found no differences in the medullary reactions between the corticolous and the 
saxicolous specimens in our area. All other differences are small and can be ascribed to the 
difference in substrate. The rather small lobes, ascending margins, dull, emaculate upper 
cortex, chemical properties, and ecological requirements are the same (in one of our 
localities, a lava flow in Kenya at 1000 m altitude, the species was found both on shrubs and 
rock, and there was no difference between them). We therefore reduce P. mordenii to 
synonymy with P. praesorediosa. 

In our area the saxicolous form has been collected in Ethiopia, Kenya, and Uganda, the 
corticolous form in Kenya and Tanzania, both in fairly dry, well lit sites between c. 700 and 
1800 m altitude. The species is pantropical to southern temperate, with its main distribution 
area in Central America (Hale, 1965). 

East African records 

Ethiopia. Shewa Province, Wondo Gennet, Winnem 422/6, 7 (O). Sidamo Province, W of Dilla, 
E 19/4. Kenya. Eastern Province, Machakos District, lava flow 5 km NW of Kibwezi, K 20/104, 105, 
504. Tanzania. Eastern Province, Morogoro District, Kitulangalo Forest Reserve, 6 May 1978, Dahl 
(O); Morogoro District, c. 25 km N of Morogoro, on the way to Dodoma, 3 May 1978, Dahl (O); 
Morogoro District, Morogoro, above university campus, 30 April 1978, Dahl (O). Uganda. East Mengo 
District, Kyagwe County, Owen Falls Dam, facing Jinja, U 5/1-2, 10. Masaka District, Bukoto 
County, rock outcrop by road to Lake Nabugabo, 3U 23/3, 4. 



47. Parmelia pseudocrinita des Abb. 

Bull. Inst.fr. Afr. noire A, 20 : 19 (1958). — Parmotrema pseudocrinitum (des Abb.) Hale, Phytologia 
28:338 (1974). Type: Guinea, Fouta-Djalon Mountains, Dalaba, 5 February 1951, des Abbayes 
(REN — lectotype, not seen). 

Thallus corticolous, rarely saxicolous, coriaceous, adnate to loosely attached, pale grey, 
grey, or grey-green. Lobes 0*5—1 (1*5) cm broad, rounded, crenate, ciliate, cilia 1-2 mm long. 
Upper side smooth, often shiny, emaculate. Medulla white. Underside black, with a brown, 



206 H. KROG & T. D. V. SWINSCOW 

naked marginal zone, rhizines abundant, long, sometimes branched. Soralia absent. Isidia 
cylindrical, simple or branched, often ciliate, abundantly produced over most of the thallus. 
Apothecia rare, laminal, up to 5 mm in diameter, thalline exciple isidiate, disc at first 
imperforate, becoming perforate, spores 15-18 x 6-8 (10) jum. Conidia filiform, 10-12 jum 
long. 
TLC: gyrophoric acid, atranorin. 

Parmelia pseudocrinita has its main distribution area in Africa south of the Sahara. In East 
Africa we have collected it in fairly well lit sites in inselbergs, riverine and low montane 
forests, and artificial habitats such as parklands and town avenues. It was mainly found 
between c. 700 and 1800 m altitude, with the exception of one collection from 2600 m. The 
species is also known from Thailand (Kurokawa, Lich. Rar. Crit. Exs. no. 33 (O)). 

East African records 

Ethiopia. See Winnem (1975). Kenya. Coast Province, Taita District, near school W of Wundanyi, 
2K27/112. Eastern Province, Machakos District, Ol Doinyo Sapuk, 20km SE of Thika, 2K4/106; 
Meru District, near Chogoria, 3 K 7/1 12. Western Province, Kakamega District, Kakamega Forest by 
River Ikuyawa, 4K 10/108. Tanzania. Northern Province, Arusha District, Mt Meru Crater, T 5/154, 
Arusha National Park, near Ngurdoto Gate, T 6/103. Eastern Province, Morogoro District, 
Kitulangalo Forest Reserve, 6 May 1978, Dahl{0). Tanga Province, Lushoto District, Usambara Mts, 
Amani, Dodwe (road towards Derema), Vitikainen 9412, 9413 (H). Uganda. West Mengo District, 
Busiro County, 3 km SW of Funjo Church, U 2/2, 5, 10; Kyadondo County, Makerere University 
grounds, U 1/2. Ankole District, Igara County, Kalinzu Forest, 2U 16/29-1. 



48. Parmelia pseudograyana Hale 

Phvtologia 27 : 2 (1973). Type: South Africa, Natal, Drakensberg, Cathedral Peak area, Nhlonhlo 
Valley, 1 500 m, E. A. Schelpe 102 1 (BOL— holotype, not seen). 

Thallus saxicolous, firmly attached, pale grey to ash grey. Lobes 0*5—1 cm broad, with 
ascending margins, deeply crenate, ciliate, cilia 0*2—1 (2) mm long. Upper side emaculate, 
usually shiny, matt and rugose towards the centre. Medulla white. Underside black, with a 
dark brown, naked marginal zone, rhizines fairly coarse, scattered, black to brown. Soralia 
on lateral lobes, marginal and linear to submarginal and punctiform, soredia often dark grey. 
Only immature apothecia seen. Conidia rod-shaped to weakly sublageniform, 8-9 /zm long. 
TLC: fumarprotocetraric acid, protocetraric acid, atranorin. 

Parmelia pseudograyana is very close to P. grayana, and, as Hale (1973) pointed out, the 
two species are morphologically indistinguishable. The conidia are also the same. Since 
mature apothecia have not yet been found in P. pseudograyana, we have been unable to 
compare spore size, but most saxicolous Amphigymnias in our area have small spores. If we 
hesitate to reduce P. pseudograyana to synonymy with P. grayana it is because their 
diagnostic medullary substances are not closely related; furthermore, P. pseudograyana, 
although more common in East Africa, has a much more restricted distribution than P. 
grayana. 

Parmelia taitae, newly described in this work, may be regarded as the parent morph of P. 
pseudograyana. 

Parmelia pseudograyana grows on acidic rock in dry, well lit sites and in our area has been 
collected between 1400 and 1900 m altitude. It also occurs in South and West Africa, but is 
not yet known outside the African continent. 

East African records 

Ethiopia. Sidamo Province, between Shashemenne and Malge Wondo, Winnem 489/2 (O). Kenya. Rift 
Valley Province, Nakuru District, entrance of Masai Gorge N of Lake Naivasha, K 30/108. Eastern 
Province, Machakos District, Lukenia, 30 km SE of Nairobi, 3K 1/134. Coast Province, Taita District, 
vicinity of Mwanda Summit, 2K 28/106, 2K 29/101, 2K 31/103. Tanzania. [? Southern Province, 
Songea District] Lupembe, Ludeka, Eusebia (F, fide Hale (1973), not seen). Uganda. Kigezi District, 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



207 



Rubanda County, 1 km W of Hamurwa, 3U 46/3. Ankole District, Igara County, 1 km E of Lubare 
Ride Pass, Lye L 398 (herb. Lye). 



49. Parmelia rava Krog & Swinscow sp. nov. 



Fig. 23 



Thallus corticola, ravus. Lobi 0*5—1 (1*5) cm lati, integri vel leniter crenati, eciliati, superne 
emaculati. Soralia marginalia, vulgo ad laminam extensa, sorediis granulosis. Apothecia ad 
laminam, disco imperforato. Spori 18-20 (22) x 8-10 jum. Conidia filiformia, 10—12 //m longa. 
Acidum protocetraricum, acidum usnicum, et atranorinum continens. 

Thallus corticolous, adnate or loosely attached, pale yellow-grey. Lobes 0*5—1 (L5)cm 
broad, entire or weakly crenate, eciliate. Upper side emaculate, matt, more or less rugose and 
cracked towards the centre. Medulla white. Underside black, with a brown, naked marginal 
zone, rhizines short. Soralia marginal and linear on peripheral lobes, subcapitate on 
ascending lateral lobes, often spreading laminally, soredia granular, pale green-grey. 
Apothecia sparse, laminal, thalline exciple sorediate, disc imperforate, spores 
18-20 (22) x 8-10 jum. Pycnidia rare, conidia filiform, 10-12 jum long. 

TLC: Protocetraric acid, ± fatty acid, usnic acid, atranorin. 

Type: Ethiopia. Sidamo Province, 5 km NW of Zenbe Woha, open woodland, 1770 m, 10 
January 1972, coll. H. Krog no. E 12/5 (O— holotype, BM, UPS— isotypes). 

Winnem (1975) determined specimens of P. rava from Ethiopia as P. dominicana Vainio, 
another sorediate Amphigymnia species with protocetraric acid, usnic acid, and atranorin. 
However, she noted that the soralia of the Ethiopian specimens were not distinctly yellow in 
contrast to those of the type specimen. Parmelia dominicana can be distinguished from P. 
rava by its mainly grey upper cortex, yellow soralia, slightly smaller spores, and sub- 
lageniform conidia c. 6 jum long. It has not been correctly reported from Africa. 
Parmelia rava cannot be distinguished from P. dilatata in the sterile state without a 




Fig. 23 Parmelia rava Krog & Swinscow, holotype (O). Rule = 1 cm. 



208 H - KROG & T. D. V. SWINSCOW 

chemical test, but aside from a more complex chemistry, P. dilatata has larger spores, 
25-27 //m long. 

Parmelia rava is present in all four countries of our study. It has been collected on trees 
and shrubs in bushed grassland and open woodland betwen 1250 and 1800 m altitude. 
Outside our area we have seen specimens from south-west and south-east Africa. 

East African records 

Ethiopia. Sidamo Province, W of Dilla, Winnem 470/15 (O); 6 km NW of Wadera, E 10/6; 5 km NW 
of Zenbe Woha, E 12/5 (type collection). Kenya. Eastern Province, Machakos District, lava flow 5 km 
NW of Kibwezi, 3 K 23/128. Tanzania. Southern Highlands Province, Iringa District, 50 km S of Iringa 
along road to Mbeya, Bjornstad AB 1 389— h (O); Iringa District, Ruaha National Park, in Brachystegia 
woodland at Magangwe Airstrip, Bjornstad AB 1499-d (O). Uganda. Masaka District, Mawogola 
County, 5 km NW of Ntusi, Lye L596 (herb. Lye). 

Additional specimens studied 

Angola. Bie, between Coemba and Munhango, 1300-1400 m, 10 February 1960, Degelius (herb. 
Degel.). Malanje, Duque de Braganca, 1 100 m, 6 March 1960, Degelius (herb. Degel.). Moxico, Vila 
Lus., Rio Dala, 1300 m, 13 February 1960, Degelius (herb. Degel.); between Luso and Cachipoque, 
1300 m, 16 February 1960, Degelius (herb. Degel.); Lucusse, River Lunque-Bungo, 1200 m, 13 
February 1960, Degelius (herb. Degel.). Zaire. Southwest of Albertville [Kalemi], 5 March 1930, Hoeg 
(TRH). Zambia. Mt Makulu Research Station near Chilanga, Angus M 1692 (BM). Zimbabwe. South 
of Felixburg, 4 February 1930, Hoeg (TRH). Mazse District, Christon Bank, Letcher 17 (BM). 
Mozambique. Nyassa, Mt Massangulo, Sousa 12 (BM). Malawi. Ncheese Forest Reserve, Burn Davy 
1418 (BM). 



50. Parmelia reticulata Taylor 

in Mackay, Flora Hibernica 2 : 148 (1836). — Parmotrema reticulatum (Taylor) Choisy, Bull. mens. 

Soc. linn. Lyon 21 : 1 75 (1952). Type: Ireland, Kerry County, near Dunkerron (not seen). 
Parmelia pseudoreticulata Tavares, Port. Acta biol. IB: 138 (1945). — Parmotrema pseudoreticulatum 

(Tavares) Hale, Phytologia 28:338 (1974). Type: Portugal, Estremadura, Arredores de Lisboa 

(LISU — holotype, not seen). 
Parmelia simulans Hale, Phytologia 22 : 32 (1971). — Parmotrema simulans (Hale) Hale, Phytologia 

28 : 339 (1974). Based on Parmelia macrocarpoides var. subcomparata Vainio, Acta Soc. Fauna 

Flora fenn. 7 (7) : 43 (1890). Type: Vainio, Lichenes Brasilienses Exs. 918, MinasGeraes, Sitio, 1885 

(BM — isolectotype). [TLC: caperatic acid, atranorin.] 

Thallus corticolous, saxicolous, or terricolous, membranaceous to coriaceous, loosely 
attached, pale grey, ash grey, or grey-green. Lobes 0-5-1-5 (2) cm broad, in part rounded and 
deeply crenate, in part irregularly incised and laciniate, sparingly to moderately ciliate, cilia 
02-2 mm long. Upper cortex reticulately maculate and cracked. Medulla white. Underside 
black, rhizinate or papillate to the margins or with a brown, naked marginal zone, rhizines 
abundant, simple or squarrose. Soralia marginal, linear to subcapitate, more rarely sub- 
marginal and punctiform to more or less pustular. Apothecia submarginal to laminal, 
substipitate, thalline exciple sorediate, disc imperforate or narrowly perforate, spores 
1 3-1 8 x 8-1 jim. Conidia filiform, (8) 1 2-1 6//m long. 
TLC: (1) salazinic acid, atranorin; (2) caperatic acid; atranorin. 

Parmelia reticulata is a species with a wide ecological amplitude and a correspondingly wide 
morphological variation. Specimens growing in rather dry sites exposed to high light 
intensity tend to become coriaceous, have weakly developed maculae, and often submarginal 
soralia, while specimens in moist, shady habitats usually are membranaceous, have marginal 
soralia, and well developed maculae. A naked marginal zone on the underside may be broad, 
narrow, or absent. Development of the soralia, whether linear, subcapitate, or pustular, is 
not correlated with other characters and does not form a basis for a division at species level. 
We consider that P. pseudoreticulata Tavares falls well within the variation range of P. 
reticulata. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 209 

Only one of the specimens from East Africa, Lye L 3 1 , corresponded to the simulans 
chemotype. However, neither that specimen nor others, from Brazil (BM), showed morpho- 
logical characters by which to separate P. simulans from P. reticulata. We strongly suspect 
that the species pair P. cetrata — P. reticulata comprises chemotypes with caperatic acid (P. 
macrocarpoides Vainio — P. simulans) and norlobaridone (P. homotoma Nyl. — P. com- 
mensurata Hale), as well as salazinic acid, but since most of these chemotypes are absent 
from our area, we have not studied the problem in detail. 

Parmelia reticulata is extremely common and widespread in all countries of our study; it 
has probably been undercollected in Uganda. Most of the specimens grew between 1 500 and 
3000 m altitude, but there are also some from around 1000 m and one collection from sea 
level. Outside our area the species is common and widespread throughout the tropical and 
temperate regions. The chemical diversity seems to be centred in Brazil. 

Selected East African records 

Ethiopia. Shewa Province, Wondo Gennet, 20 km S of Shashemenne, E 5/20. Sidamo Province, 
Wadera, E 8/3, 12 km S of Kebre Mengist, E 13/5, 30 km SE of Hagere Selam, E 15/30, W of Dilla, 
E 19/6, Yirga Alem, Sidamo Provincial Hospital, Winnem 476/22 (O). Gemu Gofa Province, Gidole, 
E 26/6. Arussi Province, W slope of Mt Chilalo, Winnem 751/34 (O). Kenya. Central Province, Nyeri 
District, Mt Kenya, W side, Naro Moru track, K 13/107; Kiambu district, escarpment E of Rift Valley, 
35 km NW of Nairobi, K 15/107; Kirinyaga District, Mt Kenya, S side, 2 km NW of Irangi Forest 
Station, K48/1 12. Eastern Province, Machakos District, Mua Hills, 3 K 2/121; Embu District, Embu, 
Izaak Walton Inn, K 53/122; Mem District, Mt Kenya, E side, at Themwe, 3K 16/241, Mt Kenya, N 
side, Sirimon track, 4K 24/122; Marsabit District, Mt Marsabit, vicinity of Lake Paradise, 4K 6/144. 
Coast Province, Kwale District, 2 km N of Gazi, in mangrove, K 44/107; Taita District, W of 
Wundanyi, 2K 26/106. Rift Valley Province, Kajiado District, Ngong Hills, K 45/4; Elgeyo Marakwet 
District, Cherangani Hills 10 km S of Labot, 2K 8/122; Uasin Gishu District, 5 km NW of Timboroa 
summit, 2K 19/106; Samburu District, WSW slope of Warges, 4K3/118; Kericho District, Kericho 
Tea Hotel, 4K 7/108. Western Province, Kakamega District, Kakamega Forest, 0*5 km SW of Forest 
Station, 4K 9/116; Nyanza Province, Kisii District, 6 km E of Keroka, 4K 1 1/108. Tanzania. Northern 
Province, Arusha District, Arusha National Park, Kusare Forest, T 2/123. Morogoro Province, 
Morogoro District, Uluguru Mts, Morningside Research Station, Ryvarden 11572 (O). Tanga 
Province, Lushoto District, Usambara Mts, Magamba c. 4 km N of Lushoto, Ryvarden 10910 (O). 
Southern Province, Mbeya District, between Kimondo and Kikondo, 9 May 1978, Dahl (O). Uganda. 
Karamoja District, Matheniko County, southern slopes of Mt Moroto, 2U 35/10. Masaka District, 
Bukoto County, by Lake Nabugabo, 3U 20/7. Kigezi District, Bufumbira County, 1km NW of 
Rushasha, at edge of Impenetrable Forest, Lye L 3 1 (herb. Lye). 

51. Parmelia rimulosa Dodge 

Ann. Mo. bot. Gdn 46 : 133 (1959). — Parmotrema rimulosum (Dodge) Hale, Phytologia 28 : 338 
(1974). Type: [South Africa] Table Mountain, John Macgillivray s.n. (BM — holotype). [TLC: 
alectoronic acid, #-collatolic acid, skyrin, atranorin.] 

Thallus corticolous, membranaceous, loosely attached, pale grey to grey-green. Lobes 
0*5—1 5 cm broad, rounded, entire or weakly crenate, ciliate, cilia slender, 1-3 mm long, 
mainly on lateral lobes. Upper side emaculate, cortex fragile, cracking and flaking. Medulla 
white, except for scattered patches of an ochraceous, K+ purple pigment (skyrin), turning 
dark grey where the cortex has sloughed off. Underside black, with a pale brown, naked 
marginal zone, rhizines long, slender, unevenly distributed. Soralia laminal, mainly 
submarginal, pustular, producing granular soredia and corticate granules. Apothecia not 
seen. (According to Hale (1965) the apothecia are perforate, with spores 20-22 x 10-13 /m\.) 
Conidia shortly filiform, 8-10 //m long. 

TLC: alectoronic acid, tf-collatolic acid, ± skyrin, atranorin. 

Parmelia rimulosa is a species of inselbergs and montane forests, where it has been collected 
between 1800 and 2900 m altitude. Outside our area it is known only from South Africa 
(Hale, 1965). 



210 H. KROG & T. D. V. SWINSCOW 

East African records 

Kenya. Rift Valley Province, Kajiado District, Ngong Hills, K45/111; Elgeyo Marakwet District, 
Sogotio Forest 8 km N of Chebiemit, 2K 10/107; Uasin Gishu District, 5 km NW of Timboroa 
Summit, 2K 19/1 19; Laikipia District, Burguret, 4K25/125. Central Province, Nyeri District, Mt 
Kenya, W side, 2K 33/109, Aberdare Mountains, 10 km W of Tusha, 3K 31/140. Eastern Province, 
Meru district, Mt Kenya, E side, at Themwe, 3K 16/128. Tanzania. Northern Province, Arusha 
District, Mt Meru Crater, T 5/104; Arusha District, Mt Meru, confluence of River Jekukumia with 
River Ngare Nanyuki, T 11/119. 

52. Parmelia sancti-angelii Lynge 

Ark. Bot. 13 (13) : 35 (1914).— Parmotrema sancti-angelii (Lynge) Hale, Phytologia 28 : 339 (1974). 
Type: Brasiliae civit. Rio Grande do Sul, Santo Angelo pr. Cachoeira, 25 January 1893, leg. G. A. 
Malme{S — lectotype). [TLC: gyrophoric acid, atranorin.] 

Thallus corticolous or rarely saxicolous, membranaceous to coriaceous, adnate to loosely 
attached, pale green, grey-green, or pale grey. Lobes 05—1 5 cm broad, rounded, crenate, 
often deeply divided and with ascending lobe margins, cilia slender, 1-3 (4) mm long, simple 
or bifurcate. Upper side emaculate, more or less folded or rugose towards the centre. Medulla 
white. Underside black, with a brown or mottled, naked marginal zone, rhizines fairly long, 
abundant. Soralia marginal, linear, sometimes spreading submarginally, soredia farinose. 
Apothecia not seen in East African material. (According to Hale (1965) the apothecia have 
imperforate discs and spores 13-18 x 7-10 //m.) Pycnidia rare, conidia weakly sublageni- 
form, 6-8 (10) jum long. 
TLC: gyrophoric acid, atranorin. 

In East Africa P. sancti-angelii is usually membranaceous and of a distinctly green colour; 
only rarely does it become more coriaceous and grey-green to pale grey. Some of our 
specimens come morphologically very close to the type. 

Parmelia lophogena is another sorediate species in our area with a membranaceous thallus 
and gyrophoric acid in the medulla. It differs from P. sancti-angelii in having granular 
soredia often interspersed with isidia, filiform conidia, and fatty acids in addition to the 
gyrophoric acid. Parmelia indoafra, which has farinose soredia and gyrophoric acid, differs 
in a more coriaceous thallus, filiform conidia, and the presence of norlobaridone. Parmelia 
permutata differs mainly in its pigmented medulla, and P. subschimperi in its maculate 
upper cortex and filiform conidia. 

Parmelia sancti-angelii is, in our area, a species of the lower montane forest and of 
artificial habitats such as parklands, plantations, and town avenues, where it has been 
collected between 1000 and 2800 m altitude. According to Hale (1965) it is a common and 
widespread pantropical species. 

Selected East African records 

Ethiopia. See Winnem (1975). Kenya. Central Province, Kirinyaga District, Mt Kenya, S side, 2 km 
NW of Irangi Forest Station near River Ena, K 48/1 14; Nyeri District, Mt Kenya Safari Club, 
4K 19/114; Fort Hall District, Aberdare Mountains, Kimakia Forest Station Ryvarden 9028 (O). Rift 
Valley Province, Uasin Gishu District, Sclater's Road 10 km SE of Lessos, 2K 18/101. Eastern 
Province, Meru District, near Chogoria, 3K7/1 19. Western Province, Kakamega District, Kakamega 
Forest by River Ikuyawa, 4K 10/110. Tanzania. Northern Province, Arusha District, Arusha National 
Park, Juniper Hill, T 3/105, Mt Meru, E side, T 10/101; Moshi District, Kilimanjaro, E of Lemosho 
Glades, Santesson 21058 (UPS). Eastern Province, Morogoro District, above Morningside in the 
Uluguru Mountains, 6 May 1978, Dahl (O). Uganda. W Mengo District, Busiro County, Entebbe 
Botanical Garden, U 3/1 ; Kyadondo County, Makerere Hill, Burnet 232 (BM). Toro District, Burahya 
County, Fort Portal, 2U 17/9. Masaka District, Bukoto County, by Lake Nabugabo, 3U 20/12. 

53. Parmelia soyauxii Mull. Arg. 

Linnaea 9 : 32 (1880).— Parmotrema soyauxii (Mull. Arg.) Hale, Phytologia 28 : 339 (1974). Type: 
Pungo Andongo in Angola, 1 880, Soyaux 246 (G — holotype). [TLC: lecanoric acid, atranorin.] 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 21 1 

Thallus saxicolous, coriaceous, adnate, strongly attached, pale grey. Lobes rounded, 
0*5-0-8 (1) cm broad, eciliate, more or less imbricate. Upper side emaculate, shiny, matt and 
somewhat rugose towards the centre, sometimes with secondary lobules. Medulla white. 
Underside black, with a brown or mottled, naked marginal zone, rhizines short, coarse, 
sparse. Soralia and isidia absent. Apothecia common, submarginal, stipitate, up to 1 cm 
broad with age, thalline margin involute, disc perforate, spores 10-13 x 5-8 jum. Conidia 
filiform, 12-16 (18)/*mlong. 
TLC: lecanoric acid, atranorin. 

Parmelia soyauxii may be considered the parent morph of P. defecta with which it some- 
times grows. In East Africa it has been collected on acidic rock fully exposed to the sun, 
between 1000 and 2100m altitude. It is also known from West and South Africa and 
Madagascar (Hale, 1965). 

East African records 

Kenya. Rift Valley Province, Uasin Gishu District, rock outcrop near Sergoi, 27 km S of Chebiemit, 
2K 9/103. Eastern Province, Machakos District, Lukenia, 30 km SE of Nairobi, 3K 1/1 18, lava flow 
5 km NW of Kibwezi, K 20/24. Coast Province, Taita District, N of Mwanda Summit, 2K 28/104. 
Tanzania. Lake Province, Bukoba District, Bukoba, gegen Uganda, Stuhlmann 405 g (G — holotype of 
P. nitens Mull. Arg.). Uganda. Kigezi District, Rubanda County, 1 km W of Hamurwa, 3U 46/202. 
Bunyoro District, Bugangaizi County, 18 km NE of Kakumiro, 3U 67/5. Masaka District, near Towa 
Forest, Sese, Thomas Th 3028 (BM). 



54. Parmelia stuhlmannii Dodge 

Ann. Mo. bot. Gdn 46 : 137 (1959). Based on Parmelia nitens f. isidiosa Mull. Arg., Bot. Jb. 20 : 255 
(1894). Type: [? Tanzania, Seengebiet] Ririre Karapo, Stuhlmann 3301 (G — holotype). [TLC: 
lecanoric acid, atranorin.] 

Thallus saxicolous, adnate, strongly attached, pale grey. Lobes 0'6-0*8 cm broad, rounded, 
entire or crenate, eciliate, more or less imbricate in central parts. Upper side emaculate, 
shiny. Medulla white. Underside black, with a brown or mottled, often matt marginal zone, 
rhizines fairly coarse. Isidia laminal, coarse. Apothecia submarginal, substipitate, thalline 
exciple smooth or isidiate, disc usually perforate at an early stage, spores 11—13 x 6-7 jum. 
Conidia filiform, 12-16 (18) jum long. 
TLC: lecanoric acid, atranorin. 

Parmelia stuhlmannii resembles P. tinctorum in most important characters, such as small 
spores, filiform conidia, and the content of lecanoric acid; furthermore, coarse isidia are 
present in the pseudotinctorum morphotype of P. tinctorum as here interpreted. However, P. 
stuhlmannii differs in its smaller lobes, strongly attached thallus, and apothecial perforations 
which are present even in young apothecia; it is always saxicolous, and does not show the 
variation in vegatative propagules as found in P. tinctorum. It is obviously closely related to 
the primary species P. soyauxii. Unfortunately, the type of P. stuhlmannii is small and 
scrappy, and it cannot be entirely ruled out that it might represent P. tinctorum. However, in 
our interpretation of the species we rely on the original description in which it was 
considered to be a form of P. nitens Mull. Arg. (=P. soyauxii). 

The main character which distinguishes this species from P. tinctorum in the field is its 
strongly attached thallus which makes it difficult to collect without a piece of the rock on 
which it is growing. An attempt at scraping it off the substrate with a knife usually results in a 
total fragmentation of the thallus. Saxicolous specimens of P. tinctorum, on the other hand, 
are always easily collected. In fertile specimens of P. stuhlmannii the presence of perfora- 
tions, even in young, cupuliform apothecia, is an added character. 

Parmelia stuhlmannii has been collected on acidic rock in dry, well lit sites between 1 500 
and 2 1 00 m altitude. It is not known outside Africa. 



212 



H. KROG & T. D. V. SWINSCOW 



East African records 

Kenya. Rift Valley Province, Uasin Gishu District, near Sergoi, 27 km S of Chebiemit, 2K 9/9; Nakuru 
District, 5 km WNW of Gilgil, 5K 18/8. Coast Province, Taita District, N of Mwanda Summit, 
2K28/104a. [? Tanzania] Ririre Karapo, Stuhlmann 3301 (G — type collection). Uganda. Ankole 
District, Nyabushozi County, 20 km E of Mbarara, near county boundary, 2U 22/12. 



55. Parmelia subarnoldii des Abb. 



Fig. 24 



Mem. Inst, scient. Madagascar B, 10: 113 (1961). — Parmotrema subarnoldii (des Abb.) Hale, 
Phytologia 28 : 339 (1974). Type: Madagascar, Centrum medium, in silva dicta Manjakatompo, in 
montibus Ankaratra, ad truncum, 2000 m, 30 July 1956, leg. des Abbayes (US — isolectotype). [TLC: 
protocetraric acid, protolichesterinic acid, atranorin.] 
Parmelia deflectens Kurok., in S. Kurokawa (Ed.), Studies on Cryptogams of Papua New Guinea: 130 
(1979). Type: Papua New Guinea, Morobe District, Bulolo, 780 m, S. Kurokawa 5810 (TNS— 
holotype). [TLC: protocetraric acid, alectoronic acid, atranorin.] 

Thallus corticolous, membranaceous to coriaceous, loosely attached, pale grey to grey- 
green, here and there black discoloured. Lobes 0*5—1 (2) cm broad, rounded or irregularly 
incised, entire or crenate, ciliate, cilia sparse to abundant, slender, 2-\ (6) mm long. Upper 
side shiny, emaculate or faintly maculate, often uneven and pitted. Medulla white, with 
occasional patches of an ochraceous, K+ purple pigment (skyrin). Underside black, with a 




Fig. 24 Parmelia subarnoldii des Abb., 5K 5/7b (BM). Rule = 1 cm. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 213 

brown or mottled, naked marginal zone, rhizines long, simple or branched, in scattered 
groups. Soralia strictly marginal, linear on broad lobes or subcapitate on narrow, lateral 
lobes. Apothecia rare, stipitate, thalline margin dentate-lobulate or sorediate, disc 
imperforate, spores (23) 26-32 x 12-15 jum, thick- walled. Pycnidia rare, conidia sublageni- 
form, 7-8 jum long. 

TLC: (1) protocetraric acid, ± protolichesterinic acid, ± skyrin, atranorin, (2) proto- 
cetraric acid, alectoronic acid, ± skyrin, atranorin, (3) protocetraric acid, or-collatolic acid, 
± skyrin, atranorin (rare). 

Parmelia subarnoldii represents the sorediate counterpart of P. amaniensis, which has the 
same chemical variation (Table 12). The alectoronic acid strain is most common, while the 
#-collatolic acid strain is known from only two localities, one in Kenya and one in Uganda. 

The species shows great variation in thickness of thallus and length of cilia, but this seems 
mainly to be a response to the environment. A membranaceous thallus and long cilia are 
usually associated with shady, moist habitats. 

Parmelia subarnoldii grows in fairly well lit sites in coastal hills, inselbergs, and montane 
forests, between c. 500 and 3000 m altitude, but is most common between 1400 and 2400 m. 
Outside our area it occurs in Malawi and Madagascar (O), and Papua New Guinea (TNS). It 
has also been recorded from Java, Mexico, and Brazil (Hale, 1965), but with new chemical 
information these records need verification. 

Selected East African records 

Kenya. Central Province, Nyeri District, Mt Kenya, W side, Naro Moru track, K 32/107, Aberdare 
Mts, near Karuru-Gura waterfalls, 3 K 32/101; Kirinyaga District, Mt Kenya, S side, Irangi Forest 
Station, 5K 4/9b; Embu District, Embu, by Izaak Walton Inn, 3K 6/104. Rift Valley Province, Elgeyo 
Marakwet District, Cherangani Hills 10 km S of Labot, 2K 8/137. Eastern Province, Meru District, Mt 
Kenya, E side, near Chogoria, 3K 7/108, at Themwe, 3K 16/156; Marsabit District, Mt Marsabit, 
vicinity of Lake Paradise, 4K 6/140. Coast Province, Kwale District, Shimba Hills, 25 km SW of 
Mombasa, Makadara Forest, K 43/105. Tanzania. Northern Province, Arusha District, Mt Meru 
Crater, T 5/129; Moshi District, Kilimanjaro, Maua, Sitari 670 (TUR). Tanga Province, Lushoto/ 
Tanga District, Usambara Mts, Hoist 2648 (ZT). Uganda. Masaka District, Bukoto County, near Lake 
Nabugabo, 3U 21/2B, near Kasaka in Jubiya Forest, Lye L 607 (herb. Lye); Buddu County, 2 km NW 
of Bale, Lake Nabugabo, Lye L 1 38 A (herb. Lye). 

56. Parmelia subcolorata Hale 

Contr. U.S. natn. Herb. 36 (5) : 340 (1965). — Parmotrema subcoloratum (Hale) Hale, Phytologia 
28 : 339 (1974). Type: Kenya, Nyanza, Kisumu-Londiani, Tinderet Forest Reserve, 1 August 1949, 
R. A. Maas Geesteranus 5339 (LD, US — isotypes). [TLC: gyrophoric acid, pigment, atranorin.] 

Thallus corticolous, loosely attached, pale grey-green. Lobes 1-2 cm broad, crenate, ciliate, 
cilia 0*5-2 mm long. Upper side emaculate to faintly maculate, distinctly maculate in the 
vicinity of apothecia, irregularly cracked in central parts. Medulla ochraceous to salmon 
pink, pigment K— . Underside black with a broad, pale brown marginal zone, rhizines fairly 
coarse, simple or branched, in scattered patches. Soralia and isidia absent. Apothecia 
stipitate, thalline exciple rugose, maculate, thalline margin weakly dentate, eciliate or rarely 
ciliate, disc perforate, spores 13-15 (18) x 8-10 jum long. Conidia filiform, 12-16 //m long. 
TLC: gyrophoric acid, pigment, atranorin. 

Parmelia subcolorata is here interpreted as the parent morph of P. permutata. The partly 
maculate upper cortex of P. subcolorata seems to be associated with the presence of 
apothecia. Chemical properties are identical, and in both species the gyrophoric acid is 
concentrated near the upper cortex, giving rise to a colour reaction with C that shades from 
red in upper parts of the medulla to vivid yellow in lower parts. 

The species has been collected in the lower montane forests from 1 700 to 2500 m altitude. 
Unlike the widespread P. permutata, it is not known outside the African continent. In 
addition to the type, Hale (1965) cited a specimen from Zaire (herb. Degel., not seen). 



214 H. KROG & T. D. V. SWINSCOW 

East African records 

Kenya. Coast Province, Taita District, W of Wundanyi, 2K 26/104. Rift Valley Province, Trans-Nzoia 
District, Mt Elgon, S of Suam River Valley to Kapchaleva Gate, Ryvarden 9367b (O); Mt Elgon, E 
side, Holm 11:2 (UPS). Nyanza Province, Kisumu-Londiani District, Tinderet Forest Reserve, Maas 
Geesteranus (type collection). 



57. Parmelia subisidiosa (Mull. Arg.) Dodge 

Ann. Mo. hot. Gdn 46 : 87 (1959). — Parmelia cetrata var. subisidiosa Mull. Arg., Bot. Jb. 20 : 256 
(1894). — Parmotrema subisidiosum (Mull. Arg.) Hale, Phytologia 28 : 339 (1974). Type: [Tanzania] 
Usambara, Bumba, 1894, Hoist 8772 p. p. (G — holotype). [TLC: salazinic acid, atranorin.] 

Thallus corticolous or saxicolous, adnate to loosely attached, pale grey, medium grey, or 
grey-green. Lobes 0*5-1 - 2 cm broad, rounded and crenate or irregularly incised and truncate, 
ciliate, cilia 0'5-2 mm long, sometimes squarrose. Upper side reticulately maculate and 
cracked. Medulla white. Underside black, with a brown marginal zone, densely rhizinate, 
rhizines simple, bifurcate, or squarrose, extending almost to the margins. Soralia absent. 
Isidia laminal and marginal, simple or branched, often bearing cilia, rarely producing lobules 
or dissolving into granules. Apothecia rare, thalline exciple isidiate, disc imperforate (or 
perforate), spores 12-14 x 6-10 //m. Conidia filiform, 10-12 /zm long. 
TLC: salazinic acid, atranorin. 

Parmelia subisidiosa is closely related to P. cetrata and P. reticulata, and may represent the 
isidiate morph in that species group. We have collected it in montane forests and inselbergs, 
most commonly between 1800 and 2400 m altitude, with a few isolated finds up to 3200 m. 
It appears to be a widespread, pantropical to temperate species. 

Selected East African records 

Ethiopia. Sidamo Province, Gidole, Winnem 641/13 (O). Kenya. Central Province, Nyeri District, Mt 
Kenya, W side, K 37/1 14, 2K 32/103, Aberdare Mts, SE of Lesatima, 5K 13/3; Kirinyaga District, Mt 
Kenya, S side, vicinity of Irangi Forest Station, K 48/ 121, Thiba Fishing Camp, K 52/108. Rift Valley 
Province, Kajiado District, Ngong Hills, K 45/1 13; Nakuru District, Londiani Forest SE of Mau 
Summit, 2K5/1; Kericho District, Kericho Tea Hotel, 4K 15/105. Eastern Province, Meru District, 
Mt Kenya, E side, at Themwe, 3K 16/68, 127. Tanzania. Northern Province, Arusha District, Mt Meru 
Crater, T 5/139, Arusha National Park, rim of Ngurdoto Crater, T 7/105; Moshi District, Kilimanjaro, 
Mweka, Sitari 289 (TUR). Uganda. Ankole District, Igara County, Lubare Ridge, 10 km S of Rubirizi, 
2U 3/4. 

58. Parmelia subschimperi Hale 

Phytologia 23:345 (1972).— Parmotrema subschimperi (Hale) Hale, Phytologia 28:339 (1974). 
Type: Kenya, Nanyuki District, Central Province, bamboo zone, National Park road, W slope, Mt 
Kenya, 2700-3 100 m, coll. R. Santesson 22 1 50 (UPS— holotype). [TLC: norlobaridone, atranorin.] 

Parmelia balensis Winnem, Norw. J. Bot. 22: 151 (1975). — Parmotrema balense (Winnem) Hale, 
Mycotaxon 5 : 436 (1977). Type: Ethiopia, Bale Province, mountain pass between Adaba and Goba, 
3500-3600 m, coll. Winnem 355/28 (O — holotype). [TLC: gyrophoric acid, norlobaridone, 
atranorin.] 

Parmelia neolobulascens Winnem, Norw. J. Bot. 22:157 (1975). — Parmotrema neolobulascens 
(Winnem) Hale, Mycotaxon 5 : 440 (1977). Type: Ethiopia, Semen, Begemder, between Geech and 
Djinn Bahr, 3400 m, June 1971, coll. Tapper 75 (BM— holotype). [TLC: gyrophoric acid, 
atranorin.] 

Parmotrema subcompositum Hale, Mycotaxon 5:440 (1977). Type: Tanzania, Kilimanjaro 
[Northern] Province, west slope of Mt Kilimanjaro, east of Lemosho Glades, 2500-2800 m, 14 
January 1970, coll. R. Santesson 21262 (UPS— holotype). [TLC: gyrophoric acid, norstictic acid, 
atranorin.] 

Thallus corticolous or saxicolous, membranaceous to coriaceous, loosely attached, pale grey 
to grey-green, margins sometimes tinged with brown. Lobes 1-2 (3) cm broad, rounded, 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 215 

entire or crenate, ciliate, cilia 05-3 mm long, simple or bifurcate. Upper side distinctly 
maculate, more or less cracked centrally, marginal and laminal lobules occasional. Medulla 
white. Underside black, with a brown or rarely mottled, naked marginal zone. Rhizines 
abundant, unevenly distributed. Soralia marginal and submarginal on revolute lobes. 
Apothecia rare, stipitate or substipitate, thalline exciple sorediate, disc mainly perforate, 
spores 20-25 (27) x 12-15 (17) //m, relatively thick-walled. Conidia filiform, 12-16 (20) pm 
long. 

TLC: (1) gyrophoric acid, ± norstictic acid, atranorin; (2) norlobaridone, ± norstictic 
acid, atranorin; (3) gyrophoric acid, norlobaridone, ± norstictic acid, atranorin. 

The wide ecological tolerance of P. subschimperi, resulting in considerable morphological 
variation in response to the environment, and the great chemical variation have caused the 
description of several superfluous species in recent years, here reduced to synonymy. The 
situation parallels that of its parent morph, P. euneta (Table 1 1). 

The species grows to considerable size. In the alpine zone on Mt Kenya we collected a 
saxicolous specimen measuring nearly 40 cm in diameter. 

Parmelia subschimperi is a species of the montane forests and the alpine zone, where we 
have collected it between 1 800 and 3600 m altitude. It is not known outside East Africa. 

Selected East African records 

Ethiopia. See Winnem (1975) under P. balensis pro parte (excluding P. indoafra), P. neolobulascens, 
P. subcomposita, and P. subschimperi. Kenya. Central Province, Nyeri District, Mt Kenya, W side, 
Naro Moru track, several localities, Aberdare Mts, several localities; Kirinyaga District, Mt Kenya, S 
side, Irangi Forest Station, 5K4/3. Rift Valley Province, Kajiado District, Ngong Hills, K 45/1 15; 
Elgeyo Marakwet District, Cherangani Hills, 10-20 km S of Labot, 2K 7/18, 2K 8/132; Uasin Gishu 
District, 5 km NW of Timboroa Summit, 2K 19/109, 1 13; Nakuru District, 1 km S of Mau Narok, 
4K 29/107; Laikipia District, Burguret, 4K 25/127. Eastern Province, Meru District, Mt Kenya, E 
side, at Themwe, 3K 16/194, Mt Kenya, N side, Sirimon track, several localities. Tanzania. Northern 
Province, Arusha District, Mt Meru, several localities, Arusha National Park, near Ngurdoto Gate, 
T 6/102; Moshi District, Kilimanjaro, W slope, E of Lemosho Glades, Santesson 21299a (UPS). 
Uganda. Kigezi District, Bufumbira County, Muhavura, N side, U 16/11. Bugisu District, N Bugisu 
County, by Sasa Hut on Mt Elgon, 2U 44/36, 0*5 km NW of ford over Sasa River, 2U 45/14. 



59. Parmelia subsumpta Nyl. 

Flora, Jena 52: 117 (1869). — Parmotrema subsumptum (Nyl.) Hale, Mycotaxon 5:434 (1977). 

Type: Brasilia, Glaziou 1841 (H-NYL 35451 — holotype). [TLC: salazinic acid, undetermined fatty 

acid, atranorin (+).] 
Parmelia leucosemotheta Hue, Nouv. Archs Mus. Hist, nat., Paris IV, 1 : 192 (1899). — Parmotrema 

leucosemothetum (Hue) Hale, Phytologia 28 : 337 (1974). Type: Mexique, Abredores de san Luis de 

Potosi, leg. Paul Maury (PC — holotype). [TLC: salazinic acid, atranorin (+).] 
Parmotrema conferendum Hale, Mycotaxon 5 : 433 (1977). Type: Venezuela, Estado de Merida, El 

Valle, north of Merida, leg. M. E. Hale 43,291 (US — holotype). [TLC: norlobaridone, loxodin, 

atranorin (+).] 

Thallus corticolous or saxicolous, coriaceous, loosely attached, grey to pale green. Lobes 
0-5-1-5(2) cm broad, crenate, ciliate, cilia 0*2-2 (3) mm long, coarse or slender, often 
sparingly developed. Upper side shiny, strongly maculate, cortex irregularly cracked over 
most of the surface. Medulla white. Underside dark brown to black in the centre, with a 
broad, brown, mottled, or ivory marginal zone, or underside uniformly pale brown, rhizines 
dimorphous, in part long, coarse, and penicillate, in part short and slender, covering the 
lower cortex almost to the lobe margins but sometimes leaving a naked marginal zone. 
Soralia marginal, linear, becoming crescent-shaped to subcapitate, occasionally spreading on 
to the lamina on revolute lobes. Apothecia not seen in East African material. (The holotype 
specimen has perforate apothecia which, according to Winnem (1975), have spores 
13x8 jum.) Conidia filiform, (10) 12-15 jum long. 
TLC: ( 1 ) salazinic acid, atranorin (+); (2) norlobaridone, loxodin, atranorin (+). 



216 H. KROG & T. D. V. SWINSCOW 

Parmelia subsumpta belongs to a group of species with a chemical variation involving 
salazinic acid and norlobaridone (see above under 'Circumscription of the species'); its 
presumed parent morph is P. erubescens (see Table 13). We have found no morphological 
variation which is correlated with the chemical diversity. 

Parmelia leucosemotheta was believed to differ from P. subsumpta in its black underside 
with an erhizinate marginal zone (Hale, 1965). However, the type specimen has a very wide 
brown zone on the underside, and can be said to be black only in the very centre of the 
thallus. On the other hand, it does not clearly show the dimorphism of the rhizines in that 
the short, thin ones, which usually extend to the margins, are largely lacking. Among our 
specimens from East Africa there is considerable variation in the colour of the underside, 
and on the same thallus some lobes may be rhizinate or papillate to the margins, while others 
have a bare marginal zone. The thin, short rhizines are sometimes poorly developed, 
especially on specimens which have grown on rock, but this trait is not correlated with the 
colour of the underside. We have found that the various characters intergrade to an extent 
where a separation into two distinct species becomes impossible. 

Only one of our specimens, from Ethiopia (Winnem, 1975, as P. confer endd), belonged to 
the norlobaridone strain, while the rest belonged to the typical strain. The strain corres- 
ponding to Parmotrema reitzii Hale, with both salazinic acid and norlobaridone, has not 
yet been found in Africa. 

In our area Parmelia subsumpta is a species of the lower montane forest where we have 
collected it on tree trunks and larger branches, more rarely on rock, between 1400 and 
2100 m altitude. It has probably been overlooked in Uganda. Outside our area it has been 
recorded from Zaire, Transvaal, Natal, and from the Americas (Hale, 1965). 

East African records 

Ethiopia. Sidamo Province, Yirga Alem, Sidamo Provincial Hospital, E 20/20. For further records, see 
Winnem (1975). Kenya. Western Province, Kakamega District, Kakamega Forest near forest station, 
Santesson 21697 (UPS). Central Province, Kirinyaga District, Mt Kenya, S side, 2 km NW of Irangi 
Forest Station, near River Ena, K 48/120, near Castle Forest Station, K 49/145, 5K5/3. Eastern 
Province, Meru District, near Chogoria, 3K 7/102, Mt Kenya, E side, at Themwe, 3K 16/123, 157. 
Rift Valley Province, Samburu District, WSW slope of Warges, 4K3/116; Marsabit District, Mt 
Marsabit, vicinity of Lake Paradise, 4K 6/142; Laikipia District, Burguret, 4K 25/128. Tanzania. 
Northern Province, Arusha District, Mt Meru, forest on E side, T 10/107, Arusha National Park, 
Kusare Forest, T 2/1 1 1, Juniper Hill, T 3/104, near Ngurdoto Gate, T 6/1 12, valley by Seneto Pool, 
T 8/101. 



60. Parmelia subtinctoria Zahlbr. 

Symb. sin. 3 : 193 (1930).— Parmotrema subtinctorium (Zahlbr.) Hale, Phytologia 28 : 339 (1974). 
Type: Sanyingpan, N von Yunnanfu, 2400 m, 14 March 1914, Handel- Mazetti 5645 (WU — 
holotype). [TLC: salazinic acid, norlobaridone, atranorin (+).] 

Thallus corticolous, usually membranaceous, loosely attached, grey-green. Lobes 
05—1 .5 cm broad, rounded, crenate, ciliate, cilia 0'2-2 mm long, simple or rarely branched. 
Upper side distinctly maculate, irregularly cracked. Medulla white. Underside brown to 
black in the centre, brown marginally, densely rhizinate with a mixture of short and long 
rhizines, rhizines and papillae often extending to the lobe margins. Soralia absent. Isidia 
abundant, laminal, thin, simple or branched, rarely ciliate. Apothecia and pycnidia not 
found in East African material. (According to Hale (1965) the apothecia have imperforate 
discs and spores 8-1 1 x 5-8 jum.) 

TLC: salazinic acid, norlobaridone, atranorin (+). 

Parmelia subtinctoria is closely related to the sorediate P. subsumpta (see above), and we 
expect that it has the same chemical variation (Table 13). We believe that P. haitiensis Hale, 
with only norlobaridone, is conspecific with it, but we have not studied the type of that 
species. 



PARMELIA (AMPHIGYMN1A) IN EAST AFRICA 217 

In East Africa P. subtinctoria is a species of well lit sites in the lower montane forest, 
inselbergs, open hillsides, and roadside trees and gardens between 1300 and 2300 m altitude. 
It has probably been overlooked in Uganda. Its world distribution has been mapped by Hale 
(1965, Fig. 24), and it has since been recorded from the Canary Islands by 0sthagen & Krog 
(1976); it is a common and widespread species of tropical and temperate regions. 

Selected East African records 

Ethiopia. See Winnem (1975). Kenya. Central Province, Nyeri District, Mt Kenya, W side, Naro Moru 
track, K 32/1 14, Mt Kenya Safari Club, 4K 19/131; Kirinyaga District, Mt Kenya, S side, Thiba 
Fishing Camp, K 52/107. Rift Valley Province, Narok District, Enabilibil, 22 km S of Siape bridge, 
4K 31/106; Kericho District, Kericho Tea Hotel, 4K 16/108; Laikipia District, Naro Moru River 
Lodge, 2K 34/104, Thomson's Falls, Santesson 21970 (UPS); Elgeyo Marakwet District, Sogotio Forest 
8 km N of Chebiemit, 2K 10/109; Kajiado District, Ngong Hills, K 45/1 14; Nakuru District, Londiani 
Forest SE of Mau Summit, 2K 5/109; Trans Nzoia District, E of Moiben, near Hoey's Bridge, 
2K 12/1 11. Eastern Province, Machakos District, Ol Doinyo Sapuk, 20 km SE of Thika, 2K 2/107; 
Meru District, Mt Kenya, E side, at Themwe, 3K 16/178; Marsabit District, Mt Marsabit, vicinity of 
Lake Paradise, 4K 6/141. Western Province, Kakamega District, Kakamega Forest, 05 km SW of 
forest station, 4K9/114. Nyanza Province, Kisii District, 6 km E of Keroka, 4K 11/1 10. Coast 
Province, Taita District, near school W of Wundanyi, 2K 27/1 17. Tanzania. Northern Province, 
Arusha District, Arusha National Park, Juniper Hill; Moshi District, Kilimanjaro, Lyamungu, Coffee 
Research Station, Bigger 1753 (BM). 

61. Parmelia sulphurata Nees & Flotow 

Linnaea 9 : (1835). — Parmotrema sulphuratum (Nees & Flotow) Hale, Phytologia 28 : 339 (1974). 
Type: Cuba, Wright 72 (UPS — neotype, BM — isotype). [TLC: vulpinic acid, atranorin.j 

Thallus corticolous, loosely attached, grey-green. Lobes 0*5-1 cm broad, rounded, ciliate, 
cilia 0*5-1 mm long. Upper side emaculate, irregularly cracked, exposing the pigmented 
medulla. Medulla intensely sulphur yellow (K— ) or orange (K+ purple, skyrin). Underside 
black, with a mottled, naked marginal zone, rhizines long, sparse, scattered. Soralia absent. 
Isidia thin, branched, concolorous with the thallus. Apothecia and conidia not found in East 
African material. (Conidia have been seen in a specimen from Brazil, leg. Malme (O); they 
were sublageniform, c. 5//m long.) 
TLC: vulpinic acid, ± skyrin, atranorin. 

We have collected P. sulphurata only once in East Africa, on a fallen branch in the low 
coastal hills of Kenya, at 480 m altitude. Its world distribution has been mapped by Hale 
(1965, Fig. 22). Although largely pantropical, it appears to be very rare in East Africa. 

East African record 

Kenya. Coast Province, Kwale District, Shimba Hills, 25 km SW of Mombasa, Makadara Forest, 

K 43/103. 

62. Parmelia taitae Krog & Swinscow sp. nov. Fig. 25 

Thallus saxicola, laxe affixus, griseus. Lobi 1-2 cm lati, orbiculares, marginibus ascendentibus, 
crenatis, ciliis 0*5-2 mm longis, superne emaculati. Soralia isidiaque nulla. Apothecia submar- 
ginalia, ad stipites inflatos, disco ad 2 cm diametro, imperforato. Spori 12-15 (17)x 10 /zm. Conidia 
sublageniformia, 6-8 //m longa. Acidum fumarprotocetraricum, acidum protocetraricum, et 
atranorinum continens. 

Thallus saxicolous, loosely attached, pale grey. Lobes 1-2 cm broad, rounded, lobe margins 
ascending, crenate, sparingly ciliate, cilia 0*5-2 mm long. Upper side emaculate, slightly 
shiny. Medulla white. Underside black, with a dark brown, naked marginal zone, rhizines 
coarse, in scattered groups. Soralia and isidia absent. Apothecia submarginal, stipitate on 
inflated stipes, thalline exciple smooth, thalline margin dentate, disc up to 2 cm in diameter, 
imperforate, spores 12-1 5 (17) x 10 //m. Conidia sublageniform, 6-8 jum long. 
TLC: fumarprotocetraric acid, protocetraric acid, atranorin. 



218 



H. KROG & T. D. V. SWINSCOW 




Fig. 25 Parmelia taitae Krog & Swinscow, holotype (O). Rule = 1 cm. 



Type: Kenya, Coast Province, Taita District, Taita Hills, Mt Iyale, 3° 23' S, 38° 20' E, alt. 
1980 m, epilithic on granitic rock, 9 December 1973, coll. /. Nordal no. INB 727 (O— 
holotype). 

Parmelia eborina Hale, described from Central America, has similar spores and conidia but 
lacks cilia, has subsessile apothecia, and produces protocetraric acid without the addition of 
fumarprotocetraric acid. 

It is possible that P. taitae represents the parent morph of P. pseudo gray ana, a sorediate 
species which occurs in the same general area. They agree in most important characters, but 
P. pseudograyana is a somewhat smaller species with lobes rarely exceeding 1 cm in width. 

Parmelia taitae is known only from the type collection and an additional collection, 
Nordal INB 728, from a nearby locality at 2050 m altitude. 



63. Parmelia tinctorum Despr. ex Nyl. 

Flora, Jena 55 : 547 (\S72).—Parmotrema tinctorum (Despr. ex Nyl.) Hale, Phytologia 28 : 339 
(1974). Type: Canary Islands, Despreaux (?) (H-NYL 35365— holotype). [TLC: not tested.] 
Parmelia pseudotinctorum des Abb., Bull. Inst. fr. Afr. noire A, 13:973 (1951). — Parmotrema 
pseudotinctorum (des Abb.) Hale, Phytologia 28 : 338 (1974). Type: Mt Tonkoui, cercle de Man, 
Ivory Coast, 14 August 1948, des Abbayes (US — isotype). [TLC: lecanoric acid, atranorin.] 

Thallus corticolous or more rarely saxicolous, membranaceous to coriaceous, loosely 
attached, pale grey to grey-green. Lobes 1-2 cm broad, rounded, entire or crenate, eciliate. 
Upper side emaculate, shiny, becoming dull towards the centre, cortex sometimes cracking 
and flaking. Medulla white. Underside black, with a broad, brown, naked marginal zone, 
rhizines fairly coarse, in scattered groups. Isidia sparse to abundant, laminal, confluent or in 
scattered groups, brown tipped or concolorous with the thallus, simple or branched, thin and 
cylindrical or coarse and irregularly inflated, often interspersed with lobules, sometimes 
partly or wholly dissolved into granular soredia. Apothecia laminal, substipitate, thalline 
exciple isidiate, disc dark brown, imperforate or with a small perforation, often radially split, 
spores 1 3-1 5 x 7-8 pm. Conidia filiform, 12-16 pm long. 
TLC: lecanoric acid, atranorin. 



PARMELIA {AMPHIGYMNIA) IN EAST AFRICA 



219 



As mentioned under 'Circumscription of the species' we have found the variation in the 
vegetative propagules unsuitable for a taxonomic division at species level in this case. There 
is no difference between P. tinctorum and P. pseudotinctorum in characters of spores and 
conidia, perforation of the apothecia, or substrate preference. Both morphotypes have their 
main distribution between 1000 and 2000 m altitude, with occasional records up to 2700 m, 
but only P. tinctorum s. str. has been found near the coast. None of them occurs in the dry, 
low lying savanna between c. 300 and 900 m. 

Parmelia tinctorum is found in mangroves, low coastal hills, and open, well lit upland 
habitats. It is a common and widespread species of tropical and temperate regions. 

Selected East African records 

Ethiopia. See Winnem (1975) under P. tinctorum and P. pseudotinctorum. Kenya. Eastern Province, 
Machakos District, Ol Doinyo Sapuk, 20 km SE of Thika, 2K 2/108, lava flow 5 km NW of Kibwezi, 
2K22/1 12; Embu District, River Thuchi, south of Chuka, 3K 17/101; Meru District, near Chogoria, 
3K 7/106. Rift Valley Province, Kericho District, Kericho Tea Hotel, 4K 16/104; Samburu District, 
WSW slope of Warges, 4K3/112; Kajiado District, Chyulu Hills, K 39/101; Laikipia District, Naro 
Moru River Lodge, 2K 34/105; Marsabit District, Mt Marsabit, vicinity of Lake Paradise, 4K 6/143. 
Coast Province, Taita District, above Wundanyi, 2K 25/1 12; Kwale District, Shimba Hills, 25 km SW 
of Mombasa, Kivumoni Forest, K 41/103, 2 km N of Gazi, in mangrove, K 44/108, 3K 30/154; Kilifi 
District, Gedi Ruins, 3 K 25/103. Western Province, Kakamega District, Kakamega Forest, 0*5 km SW 
of forest station, 4K 9/109. Tanzania. Northern Province, Arusha District, Mt Meru Crater, T 5/152, 
Arusha National Park, Kusare Forest, T 2/106. Southern Province, Songea District, Gumbiro, Nordal 
INB 849 (O). Southern Highlands Province, Iringa District, Magangwe Ranger Post at Iriki River, 
Bjornstad AB 1825b (O). Eastern Province, Morogoro District, Morogoro, above university campus, 




Fig. 26 Parmelia tsavoensis Krog & Swinscow, isotype (O). Rule = 1 cm. 



220 H. KROG & T. D. V. SWINSCOW 

30 April 1978, Dahl(0). Uganda. E. Mengo District, Kyagwe County, Owen Falls Dam, facing Jinja, U 
5/4. W. Mengo District, Busiro County, Entebbe Botanical Garden, U9/1, 68. Kigezi District, 
Bufumbira County, near Kirwa, Wolfram mine SW of Lake Mutanda, U 22/ 15a. Karamoja District, 
Dodoth County, Kidepo Valley National Park, 3U 15/105. Ankole District, Isingiro County, Kikagati, 
on Kagera River, Burnet AMB 201 (BM). 

64. Parmelia tsavoensis Krog & Swinscow sp. nov. Fig. 26 

Thallus saxicola, adnatus, griseus. Lobi ad 5 mm lati, eciliati, marginibus integris, superne 
emaculati, laeves, nitidi. Soralia nulla. Dactyii inaperti, breviter claviformes, ad laminam loborum. 
Apothecia matura non visa. Conidia filiformia, 18-20//m longa. Acidum physodicum, acidum 
oxyphysodicum, et atranorinum continens. 

Thallus saxicolous, adnate, pale grey. Lobes up to 5 mm broad, eciliate, with entire margins. 
Upper side emaculate, smooth, shiny. Medulla white. Underside black, with a broad, brown, 
naked marginal zone, rhizines short, sparse. Soralia absent. Dactyls closed, shortly 
claviform, laminally situated towards the centre of the thallus. Mature apothecia not seen. 
Conidia filiform, 1 8-20 /im long. 

TLC: physodic acid, oxyphysodic acid, atranorin. 

Type: Kenya, Eastern Province, Machakos District, lava flow 5 km NW of Kibwezi, 
2° 35' S, 37° 51' E, on lava rock, 1000 m, January 1972, coll. H. Krog. & T. D. V. Swinscow 
no. K 20/24 (BM— holotype, O, UPS, US— isotypes). 

Parmelia tsavoensis recalls forms of P. stuhlmannii in its eciliate lobes and closed dactyls 
resembling coarse isidia, but differs in its longly filiform conidia and in chemical properties. 

The species is known only from the type locality, where it was exposed to high light 
intensity; the black volcanic scoria on which it was growing reached exceedingly high 
temperatures under the noon sun. 



65. Parmlia uberrima Hue Fig. 27 

Mem. Soc. bot. Fr. 28 : 9 (\9\6).—Parmotrema uberrimum (Hue) Hale, Phytologia 28 : 339 (1974). 
Type: Afrique equatoriale anglaise, 1912, leg. Vicomte de Poncins (PC — holotype). [TLC: 
alectoronic acid, atranorin.] 

Thallus corticolous, coriaceous, loosely attached, pale grey to green-grey. Lobes 05— 1*5 cm 
broad, rounded or irregularly incised, margins ascending, crenate, ciliate, cilia 0*5-2 mm 
long. Upper side maculate, smooth to rugose or ridged, occasionally with laminal cilia. 
Medulla white. Underside strongly rugose, black in the centre, with a very broad, white 
marginal zone, lobe-ends sometimes with small brown patches, rhizines in scattered groups, 
black, brown, or white, simple or branched, sometimes coalescing into coarse, irregular 
bundles with penicillate apices. Soralia and isidia absent. Apothecia numerous, sub- 
marginal, stipitate, up to 2 cm in diameter, thalline exciple and stipe rugose to strongly 
reticulately ridged, with coarse, ciliate, isidioid outgrowths often bearing pycnidia, thalline 
margin ciliate, dentate-lobulate, disc imperforate for a long time, now and then becoming 
perforate with age, spores (15) 17-20 x (7) 8-10 /im. Conidia shortly filiform, 10-12 jum 
long. 
TLC: alectoronic acid, atranorin. 

Parmelia uberrima resembles P. argentina Krempelh. in many respects. Both species have 
small spores, imperforate apothecia with a ciliate thalline margin, a white marginal zone on 
the underside, and only alectoronic acid in the medulla. The main distinction lies in the 
coarsely dentate thalline exciple of P. uberrima, reminiscent of that found in P. hanning- 
toniana. Conidia were extremely difficult to find in the type material of P. argentina, but 
those seen were rod-shaped and 6-8 //m long. However, they may not have been 
characteristic of the species, and observations on fresh material are needed to ascertain if this 
difference is real. If intermediates between P. uberrima and P. argentina are found, P. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



221 



(Jn/i-wi+fliet uJ*nA*ni*~ &» 




y 

Fig. 27 Parmelia uberrima Hue, holotype (PC). Rule = 1 cm. 






uberrima may be regarded as conspecific with P. argentina, but for the time being it seems 
better to regard them as distinct species. 

Parmelia uberrima is mainly a lowland species which has been collected from sea level up 
to 2000 m altitude; it was locally common on a lava flow in Kenya at 1000 m, where it grew 
on twigs and thinner branches of shrubs, accompanied by, for example, P. leonis, P. andina, 
Dirinaria spp., and Ramalina africana. It seems to prefer well lit, rather dry habitats in 
scrub, thickets, and miombo woodland. It is so far known only from East Africa. 

East African records 

Kenya. [? Eastern Province], sine loc, Poncins s.n. (PC — holotype). Eastern Province, Machakos 
District, lava flow 5 km NW of Kibwezi, K 20/121, 2K22/108, 3K23/114, 5K2/27; Kitui District, 
Mutomo, 50 km NE of Kibwezi, K21/2. Rift Valley Province, Kajiado District, Chyulu Hills, 
K 39/5 B. Coast Province, Kwale District, 2 km N of Gazi, 3K 30/134. Tanzania. Southern Highlands 
Province, Iringa District, Image Mts., NordallNB 787 (O), Eastern Province. Morogoro District, Mindu 
Forest Reserve, 4 May 1978, Dahl (O). Uganda. Masaka District, Bukoto County, 1 km E of 
Kitakomaga, Lye L705 (herb. Lye); Buddu County, near Kirumba, Lye L 225 (herb. Lye). 



66. Parmelia ultralucens Krog 

Bryologist 11 : 253 (1974). — Parmotrema ultralucens (Krog) Hale, Mycotaxon 1 : 108 (1974). Type: 
USA, Georgia, Harris Co., Dowdell's Knob, on rock, 1965, Krog s.n. (O — holotype). [TLC: salazinic 
acid, lichexanthone, atranorin.] 

Thallus corticolous or saxicolous, loosely attached, pale grey. Lobes 0"8-15cm broad, 
crenate, ciliate, cilia 0*5-2 mm long. Upper side dull, emaculate, irregularly cracked towards 
the centre. Medulla white. Underside black, with a broad, brown, naked marginal zone, 
rhizines dense, of varying length. Soralia absent. Isidia laminal, unevenly distributed, brown 
tipped, often ciliate. Apothecia and pycnidia not seen in East African specimens. (In extra- 
African material apothecia are rare, with imperforate discs and spores 1 5-1 7 x 1 0-1 2jum.) 



222 



H. KROG & T. D. V. SWINSCOW 



TLC: salazinic acid, lichexanthone (in the medulla, which reacts UV+ bright yellow), 
atranorin. 

Parmelia ultralucens is the only African Amphigymnia species which produces liche- 
xanthone in the medulla. 

The species is rare in East Africa; it has been collected twice in the Usambara Mountains 
in Tanzania at about 900 m altitude. Outside our area it is largely a pantropical to temperate 
species with its main distribution area in the southeastern US and Central America (Krog, 
1974, Fig. 2) 

East African records 

Tanzania. Tanga Province, Lushoto District, Usambara Mountains, Amani, Dodwe (road towards 

Derema), Santesson 23362 (UPS), Amani, in the surroundings of Forestry House, Santesson 23155 

(UPS). 



67. Parmelia umbrosa Krog & Swinscow sp. nov. 



Fig. 28 



Thallus corticola, griseus ad griseo-viridis. Lobi 0*5—1*5 (2) cm lati, orbiculares, integri vel crenati, 
ciliis 2-4 (5) mm longis, superne nitidi, emaculati, ad centrum irregulariter rimosi. Soralia 
marginalia, linearia, ad lobos laterales et centrales, raro submarginaliter extensa. Apothecia tantum 
immatura visa, disco perforato. Conidia filiformia, 10-14 //m longa. Acidum gyrophoricum, acidum 
protocetraricum, et atranorinum continens. 

Thallus corticolous, loosely attached, pale grey to grey-green. Lobes 0*5-1 '5 (2) cm broad, 
rounded, entire or crenate, ciliate, cilia sometimes sparingly developed on peripheral main 
lobes, usually abundant on lateral and central lobes, 2-4 (5) mm long, simple or branched. 
Upper side emaculate, irregularly cracked towards the centre. Medulla white. Underside 
black, with a broad, brown or mottled, naked marginal zone, rhizines abundant, long, 
slender. Soralia marginal, linear, on lateral and central lobes, occasionally spreading 
submarginally. Only immature apothecia seen, disc perforate. Conidia filiform, 10-14/^m 
long. 

TLC: gyrophoric acid, protocetraric acid, atranorin. 




Fig. 28 Parmelia umbrosa Krog & Swinscow, holotype (O). Rule = 1 cm. 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



223 



Type: Kenya. Eastern Province, Meru District, Mt Kenya, E side, near Chogoria, 0° 14' S, 
37° 37' E, on trees in valley, alt. 1600 m, February 1974, coll. H. Krog& T. D. V. Swinscow 
no. 3 K 7/103 (O— holotype, BM— isotype). 

Morphologically P. permutata may resemble P. umbrosa in its emaculate cortex, long cilia, 
and marginal, linear soralia, but differs in its pigmented medulla and the absence of proto- 
cetraric acid. Parmelia sancti-angelii differs from P. umbrosa in its imperforate apothecia, 
sublageniform to rod-shaped conidia, and lack of protocetraric acid. Parmelia indoafra 
differs chemically in that it has norlobaridone instead of protocetraric acid, and it also has 
less well developed cilia. 

A combination of protocetraric and gyrophoric acids is known in Amphigymnia only from 
an isotype specimen of the primary species P. blanchetii Hue in BM (Winnem, 1975). The 
type is fragmentary, and it cannot be ascertained if it is related to P. umbrosa or not. The 
protologue mentions that P. blanchetii has imperforate apothecia and cylindrical conidia 
8-9 //m long, but more material needs to be studied in order to determine whether these 
differences are real. 

Parmelia umbrosa has been collected once in Uganda at c. 1 300 m altitude and a few times 
in the lower montane forests on Mount Kenya between 1 600 and 2 100 m. Outside our area it 
occurs in Malawi (Southern Province, Mulanje District, Mulanje Mts, Lichenya Plateau, 
1 800-2000 m, Ryvarden 1 1 7 1 1 , O). 

East African records 

Kenya. Eastern Province, Meru District, Mt Kenya, E side, near Chogoria, 3 K 7/103 {type collection), 
Mt Kenya, E side, at Themwe, 3K 16/103. Central Province, Kirinyaga District, Mt Kenya, S side, 
2 km NW of Irangi Forest Station near River Ena, K 48/134. Uganda. Ankole District, Isingiro County, 
Kikagati, on Kagera River, Burnet AMB 201 (BM). 

68. Parmelia vivida Krog & Swinscow sp. nov. Fig. 29 

Thallus corticola, coriaceus, adnatus, flavo-viridis. Lobi - 5-0*8 cm lati, marginibus integris, eciliati, 
superne emaculati. Soralia isidiaque nulla. Apothecia laminalia, subsessilia, disco imperforate Spori 
20-22 x 8-10 //m. Conidia sublageniformia, 6-7 //m longa. Acidum protocetraricum et acidum 
usnicum continens. 




Fig. 29 Parmelia vivida Krog & Swinscow, holotype (O). Rule = 1 cm 



224 H. KROG & T. D. V. SWINSCOW 

Thallus corticolous, coriaceous, adnate, bright yellow-green. Lobes 0*5-0*8 cm broad, 
rounded, margins entire, eciliate. Upper side matt, emaculate, irregularly cracked towards 
the centre. Medulla white. Underside black, with a brown, naked marginal zone, rhizines 
sparse, short. Soralia and isidia absent. Apothecia laminal, subsessile, disc imperforate, 
spores 20-22 x 8-10 //m. Pycnidia numerous, conidia sublageniform, 6-7 jum long. 

TLC: protocetraric acid, usnic acid. 

Type: Tanzania, Southern Province, Songea District, Gumbiro, 10°02'S, 35°29'E, 
epiphytic in miombo woodland, 920 m, 26 December 1973, coll. /. Nordal no. INB 849 (O — 
holotype). 

We consider P. vivida to be the parent morph of P. aprica, newly described in this work. The 
species is so far known only from Tanzania. 

East African records 

Tanzania. Eastern Province, Morogoro District c. 20 km SW of Ngerengere, in miombo forest, 1 May 

1978, Dahl (O, BM). Southern Province, Songea District, Gumbiro, Nordal INB 849 (O— type 

collection). 

69. Parmelia xanthina (Mull. Arg.) Vainio 

Acta Soc. Fauna Flora fenn. 1 (7): 37 (1890). — Parmelia proboscidea var. xanthina Mull. Arg., 

Flora, Jena 67 : 616 (1884).— Parmotrema xanthinum (Mull. Arg.) Hale, Phytologia 28 : 339 (1974). 

Type: Central Madagascar, Hildebrandt s.n. (G — holotype). [TLC: caperatic acid, undetermined 

fatty acids, usnic acid, trace atranorin.] 
Parmelia nyasensis Dodge, Ann. Mo. bot. Gdn 46 : 126 (1959). Type: Mt Nchisi, Nyasaland, Brass 

16922 (NY— holotype, not seen). 
Parmelia aberrans (Vainio) des Abb., Lichenes Madagascarienses et Borbonici selecti exsiccati 

(Rennes),fasc. 1, no. 79 (1961). — Parmelia xanthina f. aberrans Vainio, Acta Soc. Fauna Flora fenn. 

7(7) : 37 (1890). Type: Brasilia, Minas Geraes, Sitio, 1000 m, ad truncum arboris, 1885, E. Vainio 

(TUR-VAIN 02758 — holotype). [TLC: gyrophoric acid, caperatic acid, undetermined fatty acids, 

usnic acid, trace atranorin.] 
Parmelia madagascariacea (Hue) des Abb., Lichenes Madagascarienses et Borbonici selecti exsiccati 

(Rennes), fasc. 1, no. 20 (1961). — Parmelia caperata var. madagascariacea Hue, Nouv. Archs Mus. 

Hist. nat.. Paris IV, 1:18 (1899). — Parmotrema madagascariaceum (Hue) Hale, Phytologia 

28 : 337 (1974). Type: Madagascar, Ambositra, 1889, Rodriguez (PC). [TLC: gyrophoric acid aff., 

caperatic acid, undetermined fatty acids, usnic acid.]. 

Thallus corticolous (or saxicolous), loosely attached, yellow-green. Lobes 0*5-1 cm broad, 
crenate, ciliate, cilia 0*3-1 mm long. Upper side dull, emaculate. Medulla white. Underside 
black, with a brown or mottled, naked marginal zone, rhizines abundant. Soredia absent. 
Isidia laminal, unevenly distributed, short, thin, brown tipped, abundantly ciliate. 
Apothecia and pycnidia not seen. (According to Hale (1965) P. aberrans has spores 
10-16 x 7-10 jum.) 
TLC: gyrophoric acid aff., caperatic acid, undetermined fatty acids, usnic acid. 

The type specimen of P. aberrans gave a C+ rose medullary reaction and undoubtedly 
contains gyrophoric acid. The East African specimens and the type of P. madagascariacea 
reacted C— , KC+ red; the corresponding spots on the TLC plates went slightly higher in 
TDA and HEF than those of a gyrophoric acid blank and the substance present in P. 
aberrans. If the negative reaction with C was due to a low concentration of gyrophoric acid, 
the spots should have been lower rather than higher on the TLC plate. Whatever the 
difference, the substances in question must be very closely related. The complex fatty acid 
pattern was the same in all specimens. It included caperatic acid, fatty acids of the reddenda 
type, and fatty acids with higher Rf values, one of which may coincide with the lower spot 
produced by protolichesterinic acid. We consider the fatty acids to be diagnostic of the 
species. The absence of gyrophoric acid or related compounds in P. xanthina is here 
interpreted as depside deficiency of no taxonomic importance. 

Hale (1965) regarded P. xanthina (including P. madagascariacea) and P. aberrans as 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



225 



distinct species on account of the reaction with C in the medulla. Almeda & Dey (1973) 
considered that all specimens with either a C+ or a KC+ reaction contained gyrophoric acid, 
and on this basis they found P. madagascariacea to be conspecific with P. aberrans. They 
believed that the name madagascariacea antedated aberrans by a year, a misconception 
which can be traced back to a printing error in Hale (1965). Des Abbayes (1958) in fact made 
the two combinations on the same page, but invalidly according to the Code (Art. 33); they 
were not validated until 1961 (see above under synonyms). [Authors who wish to maintain 
chemical species within this group should investigate the chemistry of P. nyasensis Dodge, 
which is an earlier name; the KC+ reaction cited by Hale (1965) for this species indicates that 
the name might take precedence over P. aberrans and P. madagascariacea.] 

Parmelia xanthina is a conspicuous species which is not easily overlooked; it must there- 
fore be regarded as very rare in East Africa. We have collected it once in Kenya, where it 
grew on trees on an open, well lit hillside at 1 500 m altitude. It is common and widespread in 
the Americas, and has scattered occurrences in Africa and Asia (Hale, 1965). 



East African record 

Kenya. Eastern Province, Machakos District, Ol Doinyo Sapuk, 20 km SE ofThika, 2 K 2/1 1, 109. 



70. Parmelia zollingeri Hepp 



Fig. 30 



in Zollinger, Syst. Verz. ind. Archipel 1 : 9 (1854). — Parmotrema zollingeri (Hepp) Hale, Phytologia 
28:339 (1974). Type: In litore pr. Propoli, Prov. Bantam, Zollinger 1241 (L— holotype). [TLC: 
protocetraric acid, atranorin.] 

Thallus corticolous, coriaceous, loosely attached, pale grey. Lobes 1-1*5 cm broad, rounded, 
entire or weakly crenate, eciliate (or with short cilia), central lobes more or less laciniate. 




Fig. 30 Parmelia zollingeri Hepp, K 44/ 1 6 (BM). Rule =1 cm, 



226 H. KROG & T. D. V. SWINSCOW 

Upper side emaculate, smooth, slightly rugose in older parts. Medulla white. Underside 
black, with a dark brown marginal zone, rhizines short, sparse, situated in central parts of the 
thallus. Soralia and isidia absent. Apothecia laminal, cupuliform, shortly stipitate, thalline 
exciple faintly maculate, disc imperforate, spores 18-22 (24) x 8-10 jam. Conidia sublageni- 
form, 6-8 //m long. 
TLC: protocetraric acid, ± fatty acids, atranorin. 

All specimens seen from Java, including the types of P. zollingeri, P. bogoriensis Zahlbr., 
and P. overeemii Zahlbr., had distinctly ciliate lobe margins, while none of the African 
specimens showed any tendency towards cilium production. According to Hale (1965) the 
ciliate margin is a feature mainly of Asian specimens, while those from tropical America 
lack cilia. This difference is probably of no taxonomic importance. 

In East Africa P. zollingeri has been found in a broad coastal zone between sea level and c. 
1000 m altitude, where it grew in mangroves, miombo woodland, on solitary trees in 
savannas, and at the edge of forests. It is a widespread pantropical species. 

East African records 

Kenya. Coast Province, Kwale District, 2 km N of Gazi, K 44/16, 3 K 30/140, Shimba Hills, 25 km SW 
of Mombasa, Kivumoni Forest, K41/105, K42/103, Shimba Hills, Penga Hill, Santesson 20904 
(UPS); Kilifi District, Kilifi, Nattrass 237 (BM, EA— type collection of P. litoralis Dodge). Tanzania. 
Eastern Province, Morogoro District, Nguru ya Ndege, Pocs & Mwanjabe 6483/c (herb. Pocs), Mindu 
Forest Reserve, 4 May 1978, Dahl (O), c. 20 km SW of Ngerengere, 1 May 1978, Dahl (O); Rufiji 
District, Ngulakula, Nordal INB 758 (O). Southern Province, Rungwe District, Mulinda Forest, SE of 
New Langenberg, Stolz 2577 (BM). 

71. Parmeliasp. A. 

Thallus corticolous, membranaceous, adnate, pale grey. Lobes up to 6 mm broad, rounded 
and crenate or irregularly incised, eciliate, margins entire or rarely isidiate. Upper side 
emaculate to faintly maculate, smooth, becoming rugose towards the centre. Medulla white. 
Underside black, with a broad, brown, naked marginal zone, rhizines few, very short. Isidia 
numerous, mainly laminally situated, short, slender, simple, concolorous with the thallus. 
Apothecia and pycnidia not seen. 
TLC: protocetraric acid, atranorin. 

This species comes closest to P. adspersa Vainio, classified in genus Pseudoparmelia 
(=subgenus Cyclocheila) by Hale (1976(2). Morphologically it is very similar to the lectotype 
and a syntype of that species (TUR-VAIN 02509, 02510), from the Malacca Peninsula in 
Thailand. However, these specimens, as well as an isotype ofP.filipina Hale (UPS), reduced 
by Hale (1976a) to synonymy with P. adspersa, have a much more complex chemistry. They 
contain protocetraric acid, fumarprotocetraric acid, two undetermined substances in Rf class 
5 in HEF (probably in the protocetraric acid group), and two undetermined fatty acids in Rf 
class 5 in TA. We agree with Hale in placing P. fllipina in synonymy with P. adspersa, but 
doubt whether our specimens belong in the same species. The broad, brown, naked marginal 
zone on the underside of the East African specimens, together with the sparse development 
of rhizines, indicates subgenus Amphigymnia, whereas especially the type specimen of P. 
filipina, which has subsessile, laminal apothecia with spores 10-12 x 8*5 jum, seems to come 
closer to Cyclocheila. More material, preferably fruiting, is needed before the identity of the 
East African taxon can be settled. 

East African records 

Kenya. Coast Province, Kilifi District, Gedi ruins, 3K 25/107, Mida Creek, 3K 29/1 19; Kwale District, 

2kmNofGazi,3K30/137. 

72. Parmeliasp. B Fig. 31 

Thallus corticolous, loosely attached, pale grey. Lobes 0'5-l - 5cm broad, rounded or 
irregularly incised, crenate, ciliate, cilia 0'3-2 mm long. Upper side maculate, cracked and 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



227 




Fig. 31 Parmelia sp. B, K 54/109 (O). Rule = 1 cm. 

rugose towards the centre. Medulla white. Underside entirely white, rhizines white, black- 
tipped, or black. Soralia mainly marginal, linear. Apothecia and pycnidia not seen. 
TLC: protolichesterinic acid, atranorin. 

We believe that 'species B' is the sorediate counterpart of P. leonis, newly described in this 
work. However, in the absence of both apothecia and pycnidia we feel that the relationship is 
not conclusively proved. At present it is only the entirely white underside that distinguishes 
our 'species B' from the protolichesterinic acid strain of P. hababiana. 
'Species B' grew at 1850 m altitude c. 130 km north-west of the type locality of P. leonis. 

East African record 

Kenya. Eastern Province, Machakos District, Kilima Kiu, 70 km SE of Nairobi, K 54/109. 



Acknowledgements 

We thank Dr G. Degelius, Mr K. A. Lye, Dr R. Moberg, Dr T. Pocs, Dr R. Santesson, Dr H. Sipman, 
and Mr O. Vitikainen for the loan of specimens, and the directors and curators of the herbaria BM, BP, 
EA, FH, FI, G, H, L, LD, M, PC, S, TENN, TNS, TRH, TUR, UPS, US, W, WU, and ZT for allowing 
us to study material in their care. We are particularly indebted to Dr M. E. Hale for giving us access to 
unpublished data, and for determining some of our specimens at an early stage of our studies. The 
photographs were taken by Mr B. Elgvad, the Natural History Museums, University of Oslo. H. K. 
gratefully acknowledges the receipt of grants from NORAD and from the Norwegian Research Council 
for Science and the Humanities; T.D.V.S. gratefully acknowledges the receipt of grants from the Royal 
Society and the Linnean Society of London. 



References 

Abbayes, H. des. 1958. Lichens recoltes in Guinee Francaise et en Cote d'lvoire. Bull. Inst. jr. Aft. 

noire A, 20 : 1-27. 
Almeda, F. & Dey, J. P. 1973. Chemical and nomenclatural notes on the Parmelia xanthina complex. 

Bryologistl6: 541-543. 
Cengia Sambo, M. 1937a. Lichenes Africae Orientalis Italicae. Parte I: I licheni dell' Abissinia 

meridionale e della Somalia. Nuovo G. hot. ital. II, 44 : 456^470. 
\931b. Lichenes Africae Orientalis Italicae. Parte II: I licheni dell' Eritrea e delT Abissinia 

settentrionale. Nuovo G. bot. ital. II, 44 : 47 1^483. 



228 H. KROG & T. D. V. SWINSCOW 

1938. Licheni del Kenia e del Tanganica raccolti dai rev. padri della consolata. Nuovo G. bot. ital. 



11,45:364-387. 

1940. Licheni dell' A.O.I, raccolti dal console Prof. Senni e dal centurione Ing. Giordano Ufficiali 



della milizia nazionale forestale di Addis Abeba nel 1937. Nuovo G. bot. ital. II, 46 : 437^55. 
Culberson, C. F. 1972. Improved conditions and new data for the identification of lichen products by a 

standardized thin-layer chromatographic method. J. Chromat. 72 : 1 13-125. 
Culberson, W. L. 1973. The Parmelia perforata group: Niche characteristics of chemical races, 

speciation by parallel evolution, and a new taxonomy. Bryologist 76 : 20-29. 
& Culberson, C. F. 1980. Microconidial dimorphism in the lichen genus Parmelia. Mycologia 

72: 127-135. 
Dodge, C. W. 1953. Some lichens of tropical Africa. Ann. Mo. bot. Gdn 40 : 27 1^401 . 

1959. Some lichens of tropical Africa. III. Parmeliaceae. Ann. Mo. bot. Gdn 46 : 39-193. 

Esslinger, T. L. 1977. A chemosystematic revision of the brown Parmeliae. J. Hattori bot. Lab. 

42: 1-211. 
Gliick, H. 1899. Entwurf zu einer vergleichenden Morphologie der Flechten-Spermogonien. Verh. 

naturhist.-med. Vereins Heidelberg^ : i-vi, 81-216, Tafel II, III. 
Gvelnik, V. 1932. Additamenta ad cognitionem Parmeliarum. III. Reprium nov. Spec. Regni veg. 

30 : 209-226. 
Hale, M. E. 1965. A monograph of Parmelia subgenus Amphigymnia. Contr. U.S. natn Herb. 

36(5): 193-358. 

1971a. Morden-Smithsonian expedition to Dominica: the lichens (Parmeliaceae). Smithson. 

Contr. Bot. 4 : i-ii, 1-25. 

197 \b. Two new species of Parmelia (lichens) from North America. Phytologia 22 : 92-93. 

1971c. Parmelia affluens, a new species of lichen in subgenus Amphigymnia with a yellow 



medulla. Phytologia!! : 141-142. 

— 1972. Six new species of Parmelia (Lichenes) from Africa. Phytologia 23 : 343-349. 

— 1973. New Parmeliae (Lichenes) from Africa. 2. Phytologia 27 : 1-6. 

— 1974a. Notes on species of Parmotrema (Lichenes: Parmeliaceae) containing yellow pigments. 
Mycotaxon 1 : 105-1 16. 

— 19746. New combinations in the lichen genus Parmotrema Massalongo. Phytologia 28 : 334-339. 

— 1976a. A monograph of the lichen genus Pseudoparmelia Lynge (Parmeliaceae). Smithson. Contr. 



Bot. 31 : i-iii, 1-62. 

— 19766. A monograph of the lichen genus Parmelina (Parmeliaceae). Smithson. Contr. Bot. 

33 : i-iii, 1-60. 

1977. New species in the lichen genus Parmotrema Mass. Mycotaxon 5 : 432-448. 



Henssen, A. & Jahns, H. M. 1973 ['1974']. Lichenes. Eine Einfuhrung in die Flechtenkunde. Stuttgart. 
Hue, A. 1916. Lichenes Poncinsianos in Africa tropico-orientiali anglica a Vicecomite de Poncins anno 

1912 lectos elaboravit. Bull. Soc. Bot. France 63, Mem. 6 (28) : 1-24. 
Jatta, A. 1909. Licheni del Ruwenzori. In L. A. di Savoya (Ed.), II Ruwenzori, Relazioni scientifiche 

1:1-11 (reprinted). 
Karnefelt, I. 1979. The brown fruticose species of Cetraria. Op. bot. Soc. bot. Lund 46 : 1-150. 
Klement, O. 1962. Zur Flechten-Vegetation von Tanganjika. Stuttg. Beitr. Naturk. 85 : 1-8. 
Krempelhuber, A. 1877. Neue Beitrage zu Afrika's Flechten-Flora. Linnaea 41 : 135-144. 
Krog, H. 1974. Parmelia ultralucens, a new lichen species in subgenus Amphigymnia. Bryologist 

77:253-256. 
& Swinscow, T. D. V. 1975. Variation in vegetative propagules in the African lichen Parmelia 

lophogena. BrvologistlH : 67-70. 

- 1 977. The Parmelia borreri group in East Africa. Norw. J. Bot. 24 : 1 67-1 77. 
1 979. Parmelia subgenus Hypotrachyna in East Africa. Norw. J. Bot. 26 : 1 \-43. 



Kurokawa, S. 1968. New or noteworthy species of Parmelia of Japan. J. Jap. Bot. 43 : 349-353. 

1 969. A note on some rare lichens of Japan. J. Jap. Bot. 44 : 225-229, pi. XVIII. 

1979. Enumeration of species of Parmelia in Papua New Guinea. In S. Kurokawa (Ed.), Studies 

on Cryptogams of Papua New Guinea: 125-148, Pis 1 , 2. 
Massalongo, A. B. 1860. Esame comparativo di alcuni generi di licheni. Atti Imp. Reg. 1st. Venet. Sci. 

111,5:247-276. 
Menlove, J. E. 1974. Thin-layer chromatography for the identification of lichen substances. Bull. Br. 

lich. Soc. 34 : 3-5. 
Moberg, R. 1977. The lichen genus Physcia and allied genera in Fennoscandia. Symb. bot. upsal. 

22(1): 1-108. 



PARMELIA (AMPHIG YMN1A) IN EAST AFRICA 229 

Miiller, J. 1884. Lichenologische Beitrage. XX. Flora, Jena 67 : 1-9 (Separat-Abdruck). 

1 885. Lichenologische Beitrage XXII. Flora, Jena 68 : 1-24 (Separat-Abdruck). 

1890. Lichenes Africae tropico-orientalis. Flora, Jena 73 : 334-347. 

1891. Lichenologische Beitrage. XXXV. Flora, JenalA : 371-382. 

1892. Lichenes exotici. Hedwigia 31 : 276-288. 

1893. Lichenes africani. Bot. Jb. 19 : 505-510. 

1894. Lichenes usambarenses. Bot. Jb. 20 : 238-298. 

Nylander, W. 1885. Parmeliae exoticae novae. Flora, Jena 68 : 605-6 15. 

0sthagen, H. & Krog, H. 1976. Contribution to the lichen flora of the Canary Islands. Norw. J. Bot. 

23:221-242. 
Stein, B. 1888. Ueber afrikanische Flechten. Jber. schles. Ges. vaterl. Cult. 66 : 133-142. 
Steiner, J. 1897. Flechten aus Britisch-Ostafrika. Sber. Akad. Wiss. Wien, Math.-nat. Kl. 106 : 1-28. 
Stevens, G. N. & Rogers, R. W. 1979. The macrolichen flora from the mangroves of Moreton Bay. 

Proc. R. Soc. Qd90 : 33-49, PI. 1 . 
Stirton, J. 1877-78. On certain lichens belonging to the genus Parmelia. Scott. Nat. 4:200-203, 

252-254,298-299. 
Stizenberger, E. 1890. Lichenaea africana. Fasc. I. Ber. Tat. St Gall, naturw. Ges. 1888/89 : 1-144 

(Separatabdruck). 
1891. Lichenaea africana. Fasc. II. Ber. Tat. St Gall, naturw. Ges. 1889/90:145-280 

(Separatabdruck). 

1893. Supplementa ad lichenaeam africanam. I. Ber. Tat. St Gall, naturw. Ges. 1891/92 : 86-96. 
1895. Supplementa ad lichenaeam africanam. II. Ber. Tat. St Gall, naturw. Ges. 



1893/94:215-264. 
Swinscow, T. D. V. & Krog, H. 1976. The Usnea articulata aggregate in East Africa. Norw. J. Bot. 

23:261-268. 

1979. The fruticose species of Usnea subgenus Usnea in East Africa. Lichenologist 11 : 207-252. 

Taylor, T. 1847. New lichens, principally from the herbarium of Sir William J. Hooker. J. Bot., Lond. 

6: 148-197. 
Vainio, E. A. 1890. Etude sur la classification naturelle et la morphologie des lichens du Bresil. Acta 

Soc. Fauna Flora fenn. 7 : i-xxx, 1-247. 
1898. Lichenes a G. F. Scott-Elliot in viciniis montis Ruwenzori in Africa centrali annis 1893-94 

collecti. Hedwigia (Beiblatt) 37 : 39^4. 
Vobis, G. 1980. Bau und Entwicklung der Flechten-Pycnidien und ihrer Conidien. Biblthca lich., 

Vaduz 14 :i-v, 1-141, 47 Pis. 
Winnem, B. 1975. Parmelia subgenus Amphigymnia in Ethiopia. Norw. J. Bot. 22 : 139-166. 
Zahlbruckner, A. 1909. Ergebnisse der botanische Expedition der Akademie der Wissenschaften nach 

Siidbrasilien 1901. Lichenes. Denkschr. Akad. Wiss., Wien, math.-nat. Kl. 83 : 87-21 1. 

1926. Afrikanische Flechten (Lichenes). Bot. Jb. 60 : 468-552. 

1932. Lichenes in Africa lecti. Annls Cryptog. exot. 5 : 198-275. 

& Hauman, L. 1936. Les lichens des hautes altitudes au Ruwenzori. Mem. Inst. R. colon. Beige 

5: 1-31, PL I-V. 



Index 

Specific epithets in Parmelia (subgenus Amphigymnia) and Parmotrema are listed. Accepted 
names are in roman and synonyms in italic. New names and principal references are in bold, 
whilst an asterisk (*) denotes a figure. 

aberrans 143,224,225 andina 143, 146, 148, 149, 152, 154, 159, 

abessinica 146, 148, 149, 152, 153, 154, 156, 168,169,186,192,221 

159,160,161,165,184,192 aprica 143, 148, 151, 152, 153, 163,169*, 170, 

affluens \43, 149, 177, 178 224 

aldabrensis 146, 148, 149, 151, 153, 160, araucariarum 151, 152, 1 53, 163, 170 

161,166*, 167, 191, 197 argentina 179,220,221 

amaniensis, 143, 146, 148, 149, 152, 153, arnoldii 204 

156, 157, 160, 161,167*, 168, 189,213 austrosinensis 146, 148, 151, 152, 153, 163, 

171, 176, 177 



230 



H. KROG & T. D. V. SWINSCOW 



balensis\43, 155, 188,214 

bangii 146, 151, 152, 153, 163, 171, 172, 173, 

200 
blanchetii 223 
bogoriensis 226 
breviciliata 143, 195, 196,203 

cetrata 146, 148, 149, 152, 153, 159, 172, 209, 

214 
commensurata 209 
composita 143, 155, 181, 182 
conferendum 143, 156,215 
consors 143,202 
cooperi 146, 148, 151, 152, 153, 165, 171, 

172 
corniculans 146 
crinita 146, 148, 150, 152, 153, 162, 173, 

180, 191 
cristifera 146, 151, 154, 163, 164, 173, 174*, 

175 
crvptoxantha 146, 150, 152, 153, 162, 163, 

175*, 176, 197 

dalei\ 69 

defecta 148, 151, 152, 153, 163, 171, 176*, 177, 

200,211 
deflectens 143, 156,212 
diacidula 195 
dilatata 143, 146, 149, 151, 163, 170, 177, 

178, 184,207,208 
direagens 148, 151, 152, 153, 155, 156, 164, 

178, 190 
diversa 143, 193, 197, 198 
dobs a 195 
dominicana207 
durumae 143, 146, 149, 151, 153, 159, 160, 

161,178, 179* 

eborina218 

eciliata 146,147*, 148, 149, 152, 160, 161,180 

erubescens 143, 146, 148, 149, 152, 154, 156, 

157, 160, 161,180,216 
euneta 143, 146, 147*, 148, 149, 152, 153, 

154, 155, 157, 160, 161,181*, 182, 190,215 
curvcarpa 180 
eurysaca 146, 149, 153, 160, 161,182, 183 

gardneri 146, 148, 151, 154, 163, 164, 174, 

175. 177,183*, 184 
glaucocarpa 165 
glaucocarpoides 165, 192 
grayana 146, 151, 152, 157, 164,184,206 

hababiana 146, 148, 151, 152, 156, 164, 

165,184, 185,227 
haitiensis 156, 216 
hanningtoniana 146, 149, 152, 153, 160, 161, 

185, 186*, 220 
hildebrandtii 174 



hololoba 146, 148, 149, 151, 153, 159, 160, 

161,169, 172,186, 187* 
homotoma 209 

indicum 143, 187 

indoafra 143, 151, 152, 153, 165, 187, 188*, 

210,215,223 
inexpectata 143, 146, 149, 152, 153, 157, 

160, 161,188, 189 

jacarandicola 143, 146, 148, 149, 152, 153, 
160, 161, 189*, 190 

kwalensis 143, 148, 150, 151, 153, 162, 190*, 
191 

leonis 143, 146, 149, 152, 153, 154, 159, 191*, 

221,227 
leucosemotheta 143,215,216 
litoralis 226 
lobulascens 143, 146, 148, 151, 152, 153, 

157, 164,192,193,198,204 
lophogena 150, 152, 153, 154, 162, 165, 

173,194, 197,210 
louisianae 148, 151, 152, 153, 164, 194*, 

195 

maclayana 143, 146, 147*, 148, 149, 150, 
152, 154, 160, 161, 179, 190, 195, 196* 
198,203,204 

macrocarpoides 208 

madagascariacea 143,224,225 

melanothrix 192 

mellissii 146, 150, 152, 154, 162, 163, 173, 
197 

mesotropa 147, 165 

modesta 143, 168 

mordenii 143, 205 

myelochroa 170 

natalensis 143, 192, 193 

neolobulascens 143, 155,214 

neotropicum 157, 180, 

nilgherrensis 143, 146, 148, 149, 150, 152, 

153, 154, 157, 160, 161, 193,197, 198 
nitens 196,211 
nyasensis 143, 224, 225 

over emii 226 

pachvspora 143, 156, 167, 168 
paradoxa 143, 168, 188, 189 
parahyptropa 151, 164,198*, 199 
pardii 143, 146, 151, 152, 153, 163, 199*, 200 
perforata 144 

perlata 146, 148, 151, 152, 153, 164,200 
permutata 148, 151, 152, 163, 201, 210, 213, 
223 



PARMELIA (AMPHIGYMNIA) IN EAST AFRICA 



231 



pigmentifera 143, 146, 149, 151, 153, 159, 

179,201,202* 
pilosa 143, 151, 152, 153,164,202 
planatilobata 150, 152, 153, 162,203 
poolii 148, 151, 152, 164, 165,203,204* 
praesorediosa 143, 146, 148, 151, 152, 163, 

174,205 
preperforata 167 
proboscidea 203, 224 
procera 143, 146, 195, 196 
progenes 149, 177 
pseudocrinita 146, 148, 150, 152, 153, 162, 

205, 206 
pseudograyana 148, 151, 152, 153, 157, 164, 

177,200,206,218 
pseudonilgherrensis 143, 192, 193 
pseudoreticulata 143,208 
pseudotinctorum 143, 154,211,218,219 

rampoddensis 203, 204 

rava 143, 146, 151, 152, 153, 163, 170, 207*, 

208 
recipienda 156, 180 
reitzii 156,216 
reticulata 143, 145, 146, 148, 151, 152, 153, 

163,208,214 
rigida 195 

rimulosa 146, 151, 152, 153, 163,209 
robusta 177, 184 



stuppea 174, 175 

subarnoldii 143, 146, 148, 151, 152, 156, 

164, 212*, 213 
subcaperata 157, 180, 185 
subcolorata 146, 148, 149, 152, 153, 159, 

201,213 
subcompositum 143, 155,214 
subisidiosa 146, 148, 150, 152, 162,214 
subproboscidea 179 
subrugata 168, 179 
subschimperi 143, 146, 148, 151, 152, 153, 

155, 157, 164, 188,210,214,215 
subsumpta 143, 146, 148, 151, 152, 153, 

156,164,202,215,216 
subtinctoria 146, 148, 150, 152, 156, 162, 

191,216 
sulphuratal50, 151, 162,217 

taitae 143, 146, 148, 149, 152, 153, 157, 

160,161,206,217,218* 
thomasii 1 69 
tinctorum 143, 144, 145, 146, 150, 151, 152, 

154, 162,211,218,219 
tsavoensis 143, 150, 152, 153, 162, 219*, 220 

uberrima 146, 148, 149, 151, 153, 154, 159, 

179, 192, 195,220,221* 
ultralucens 146, 150, 152, 162,221,222 
umbrosa 143, 151, 152, 153, 164, 222*, 223 



sancti-angelii 146, 151, 152, 165, 201, 210, 

223 
sieberi 1 74 
simulans 143,208 
soyauxii 146, 148, 149, 152, 153, 159, 177, 

200,210,211 
spilota 143, 155, 181, 182 
stenopteris 179 
stuhlmannii 146, 148, 150, 152, 153, 162, 

211,220 



virens 143, 180 
vivida 143, 146. 
170, 223*, 224 



148, 149, 152, 153, 159. 



wainii 179 

xanthina 143, 146, 150, 152, 162,224,225 

zollingeri 146, 148, 149, 151, 159, 184, 225*, 
226 



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Titles to be published in Volume 9 



The lichenicolous Coelomycetes. By D. L. Hawksworth 

The genus Callithamnion (Rhodophyta : Ceramiaceae) in the 
British Isles. By P. S. Dixon and J. H. Price 

Parmelia subgenus Amphigymnia (lichens) in East Africa. 

By Hildur Krog and T. D. V. Swinscow 

The genus Selaginella in tropical South America. 

By A. H. G. Alston, A. C. Jenny and J. M. Rankin 



Printed by Henry Ling Ltd, Dorchester 



Bulletin of the /f^v 

' < - ..... 

British Museum (Natural Hist " 



The genus Selaginella in tropical 
South America 

A. H. G. Alston, A. C. Jermy & 
J. M. Rankin 



Botany series Vol 9 No 4 17 December 1981 



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This number completes Volume 9 



ISSN 0068-2292 Botany series 

Vol 9 No 4 pp 233-330 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 17 December 1981 



The genus Selaginella in tropical South America 

Arthur Hugh Garfit Alston 

Anthony Clive Jermy and Josephine Margaret Rankin 

Department of Botany, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Prior to his untimely death in 1958, A. H. G. Alston had been preparing an account of the 
Selaginellas of tropical South America excluding Brazil. Following his death the manuscript 
lay fallow for a considerable period, but, such was its potential importance in providing an 
authoritative account by an established specialist of a large and critical genus from a poorly 
known major region, that my colleagues in the Museum Fern Section, Mr A. C. Jermy and 
Mr J. A. Crabbe, felt that steps should be taken to make the work available to pteridologists 
throughout the world. By this time botany had progressed, and considerable revision of the 
manuscript was essential if the publication was to be more than a fossilised version of the 
author's intentions and worthy in the 1980s of his international standing in pteridology. The 
post-Alston additions are clearly indicated in the detailed introduction that follows. I should 
like to take this opportunity of thanking my colleagues in the Fem Section — Mr A. C. 
Jermy, Mrs J. M. Rankin and Mr J. A. Crabbe (now retired), for the great amount of careful 
work that has been put into this publication. While their aim has been solely to make 
Alston's work as effective as possible for use today, the magnitude and importance of their 
own contributions, which their reticence partially obscures, should not be overlooked. 

J. F. M. Cannon 
Keeper of Botany 



Contents 

Synopsis 233 

Introduction 234 

Characters used in identification 236 

Key to species 240 

Species 250 

References 325 

Index 326 



Synopsis 

This paper completes a series of regional monographs on Selaginella written by the late A. H. G. 
Alston between 1934 and 1955. His manuscript on tropical South American species (excluding 
Brazilian, for which an account has already been published), was left in an unfinished state in the 
Department of Botany archives. It has been completed within the style of Alston's accounts, and in 
order to facilitate the identification of specimens, his account of the Brazilian species has been 
incorporated. 

133 species (and six subspecies or varieties) are keyed out and specimens seen by the authors listed 
according to geographical location. A table summarises the distribution of all taxa in the states covered 
by the monograph, namely: Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Peru, 
Surinam and Venezuela. 



Bull. Br. Mus. nat. Hist. (Bot.)9(4): 233—330 Issued 17 December 1981 

233 



234 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

The following new taxa are described and illustrated: Selaginella calceolata Jermy & Rankin, S. 
cheiromorpha Alston, S. flacca Alston, 5". meridensis Alston, S. ovifolia subsp. philipsonii Jermy & 
Rankin, S. quadrifaria Alston, S. sandwithii Alston, S. scintillata Alston, S. valdepilosa subsp. 
tricholoma Jermy & Rankin, and S. wurdackii Alston. Selaginella praestans Alston is a nom. nov. for 
S. sprucei A. Br., non Hook., whilst S. bahiensis subsp. manausensis (H.P. Bautista) Jermy & Rankin is 
a new combination. 

Somario 

Este trabajo completa una serie de monografias regionales sobre Selaginella escritas por el difunto A. 
H. G. Alston entre 1937 y 1955. Su manuscrito sobre especies tropicales sudamericanas (excluyendo 
las brasileras para las cuales un trabajo ya habia sido publicado) fue depositado, sin completarse, en los 
archivos del Departamento de Botanica. Ha sido completado al estilo de los trabajos de Alston, pero 
para facilitar la identification de especimenes su publication sobre las especies brasileras tambien ha 
sido incorporada. 

Ciento treinta y tres especies (y seis subespecies o varidades) son incluidas en la clave y los 
especimenes vistos por los autores son ordenados en listas de acuerdo con sus localidades geograficas. 
Una tabla resume la distribution de todos los taxa en los paises considerados por la monografia, siendo 
estos Bolivia, Brazil, Colombia, Ecuador, Guyana francesca, Guyana, Peru, Surinam y Venezuela. 

Los siguientes nuevos taxa son descriptos e illustrados: Selaginella calceolata Jermy & Rankin, S. 
cheiromorpha Alston, S. flacca Alston, S. meridensis Alston, S, ovifolia subsp. philipsonii Jermy & 
Rankin, S quadrifaria Alston, S. sandwithii Alston, S. scintillata Alston, S. valdepilosa subsp. 
tricholoma Jermy & Rankin, S. wurdackii Alston. Se han hechos los siguientes cambios: S. praestans 
Alston nom. nov. por S. sprucei A. Br., non Hook.; S. bahiensis subsp. manausensis (H.P. Bautista) 
Jermy & Rankin comb. nov. 

Introduction 

A. H. G. Alston was, at the time of his unfortunate and sudden death on 17 March 1958, an 
accredited specialist in the genus Selaginella. For almost 30 years he had published regional 
accounts (see Crabbe, 1960, 1969) with keys, distributional data and synonymy; but rarely, 
except when describing new species (35 in total), did he describe a taxon in any comparative 
detail. Mr Alston did not therefore communicate his vast knowledge through his printed 
works, although the herbarium at the British Museum (Natural History) was curated care- 
fully and shows Alston's ideas on relationships. Nevertheless, his concepts on classification 
within the genus or his concepts of species (he never used the rank of subspecies, but 
accepted other workers' varieties) are not at all clear. When pressed for an intra-generic 
arrangement (e.g. in Walton & Alston, 1938), he followed Baker (1887). 

At the time of his death Mr Alston was working on this paper, his last major regional 
account. He had many years before written an account of the Brazilian species (Alston, 1936) 
and the paper he was preparing was to cover those tropical countries south of Panama, 
excluding Brazil. In 1952 he published a revision of West Indies species of Selaginella 
(Alston, 1952), and three years later on those of continental North America, i.e. Panama 
northwards (Alston, 1955). For all of these works Alston studied the type specimens involved 
and it may be assumed that the types have been seen unless otherwise stated in this text. 

In 1938 he prepared an account for Pulle's Flora ofSuriname (Alston, 1938), and a year 
later published (Alston, 1939) an account of the temperate species, i.e. those in Argentina, 
Uruguay and Paraguay. It was decided that, in order to make this account as comprehensive 
as possible, the Brazilian data as published in 1936 would be included, but we must stress we 
have seen little recent material from that country, and appreciate that the distribution of 
species as shown in Table I therefore may well be changed by the recent collections of H. P. 
Bautista, A. F. and R. M. Tryon and P. Windische, amongst others. The area now covered is 
shown on Fig. 1 , namely the Guianas, Venezuela, Colombia, Ecuador, Peru, Bolivia and 
Brazil. A summary of the distribution of the species within these countries is given in Table 
1. For the revision, we have not solicited all the material of the past 20 years that must be 
available, although we have had further material sent by the curators of AAU, CGE, COL, 
IAN, K, NY, UC, US, VEN. We thank these and the Directors of B, C, E, L, M, P, S, U and 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



235 



I 

80 


1 


i i 

60 50 


i 

40 W 


v j / yO jVENEZUELA 
J COLOMBIA ) ^ 




10- 


"ECUADOR^ 






o- 


S. PERU . 








-10 \ 


^NjT BOLIVIA 


BRAZIL 




-20 








I 20- 


-30 




' 




30- 


-40 


/ 


f 

1 r 




40- 




800 KM 


-50 






50- 


\ 


so *^r" 


60 40 
i / / 


20 W 



Fig. 1 The area covered by the present account. 



236 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

UPS and Dr H. P. Bautista (INPA) for sending material when requested and for being 
tolerant over the time that some of the original material sent to Alston has been kept. Two 
papers were published (Crabbe & Jermy, 1973, 1976) in order to release loaned material of 
new species. A further seven new species are published now. All geographical names and 
provincial boundaries follow The Times Atlas of the World mid- century edition. Thus French 
Guiana records are not divided between the two provinces now maintained in that country, 
i.e. Cayenne and St Laurent du Maroni. 

We should like to record our thanks to Alison Paul (BM) for preparing the map, taking the 
SEM micrographs and compiling Table 1 and the index, to Dr N. Robson and Miss K. 
Kavanagh for help with the Latin diagnoses, to Dr S. M. Price for the translation of the 
synopsis and to J. R. Laundon for editing the manuscript with meticulous care. A special 
acknowledgement must be made to our former colleague J. A. Crabbe, who had spent much 
time in organising Alston's manuscript, notes and specimens, and who has prepared an index 
to collectors, their field numbers and the determination of their material. Due to the cost of 
production it has been decided not to publish this index but to hold it in the Fern Section 
library of the British Museum (Natural History). Researchers working on the biography of 
any botanist involved can write to the Keeper who will place any relevant data at their 
disposal. We also thank Paul Somers, Jr., for presenting us with his unpublished doctorate 
thesis (A systematic survey of the Articulatae series of the genus Selaginella and a 
monographic treatment of the S. sulcata group (sens, str.) University of Tennessee, 
Knoxville, 1978). This contains an original approach to Selaginella taxonomy. We have 
with Dr Somers's permission adopted a few nomenclatural changes clarified by him, and 
look forward to seeing the rest of his researches published in the near future. 

Characters used in identification 

Habit. Species of Selaginella range from those that are truly prostrate and wide-creeping, 
through scrambling forms eventually supported by means of 'stilt roots' (rhizophores), to 
those with various complex forms of branch patterns culminating in dendroid growth forms. 
Branching patterns are generally of two kinds. In one the whole or primary branch unit is 
finite in growth, and cones are formed synchronously at all ultimate branch apices. In the 
other type the lowermost primary branch remains sterile and continues growth only after the 
upper branches have coned and shed their spores. These two growth patterns were correlated 
with chromosome base number, the former being based on 2n = 20 and the latter on 2n = 18 
(Jermy, Jones & Colden, 1967). A further branch system is that in which the main stem is 
indefinite in growth and the primary branches are finite and often of a distinctive pattern. 
The outline of branch systems is diagnostic but may be affected by environment. Vegetative 
reproduction of dendroid species is usually by soboles or stolons. Other species, with a less 
erect 'frondose' habit and branches showing indeterminate growth, may spread by budding 
from attenuated (flagelliform) stem apices. Leaves on these extended stems or on the 
resulting young plants may not be typical in shape, size or density. 

Stem. Stem anatomy and the number of vascular bundles have been used by Hieronymus 
( 1 90 1 ) to demarcate sections within the genus. The latter character has been used in this key 
and is best examined in main stems. Stem anatomy will repay further study and has been 
reviewed by Ogura (1972) and Somers (1978). Stems may be deeply suffused with a red 
pigment (rhodoxanthin; cf. Jagels, 1969). Some species which are lightly tinged pink when 
fresh lose that colour on drying; in the key, stem colour refers to dried specimens only. Stem 
'articulations' (present in species Nos 104-133) are best seen in the dried material as a 
sunken ring or a dark zone just below the dichotomy forming a branch. Their function, if 
any, is unknown. Although the tissue organisation of the 'articulations' is not what would be 
expected of a natural abscission point, Hieronymus ( 1 90 1 ) suggests that strong winds, animals 
or the weight of the plants themselves can break the plants at these points, and Somers (1978) 
frequently observed specimens broken in this way. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 

Table 1 Distribution ofSelaginella species in the countries of tropical South America 



237 





03 
C 

'3 

a 

o 

C 
<D 
J— 


3 


03 

c 

a 
>> 

3 


_5 

N 

a 


.5 
'.5 
£ 
o 
o 


o 

03 
o 




.5 
> 


'n 
03 
t-i 




u* 


GO 


a 


> 


U 


UJ 


cu 


CQ 


03 


1 sellowii Hieron. 




















2 sartorii Hieron. 








• 








• 




3 peruviana (Milde) Hieron. 












• 


• 


• 




4 convoluta (Arnott) Spring 






• 


• 








• 


• 


5 pallescens (C. Presl) Spring 








• 












6 erythropus (Martius) Spring 




















7 umbrosa Lemaire ex Hieron. 




















8 haematodes (Kunze) Spring 




















9 coarctata Spring 


















• 


10 longissima Baker 




















11 mazaruniensis Jenman 






• 














1 2 palmiformis Alston ex Crabbe & Jermy 








• 












1 3 flabellata var. latifrons A. Braun 








• 












1 4 mortoniana Crabbe & Jermy 












• 








15 lechleri Hieron. 














• 






16 c he iromorp ha Alston 




















17 terezoana Bautista 


















• 


1 8 anceps (C. Presl) C. Presl 




















1 9 hirsuta Alston ex Crabbe & Jermy 








• 












20 bahiensis Spring subsp. bahiensis 


















• 


subsp. manausensis (Bautista) 




















Jermy & Rankin 


















• 


2 1 wolfii Sodiro 










• 


• 


• 






22 hartii Hieron. 








• 












23 praestans Alston 












• 


• 






24 amazonica Spring 








• 










• 


25 oaxacana Spring 












• 








26 quadrifaria Alston 














• 






27 tanyclada Alston ex Crabbe & Jermy 




















28 bombycina Spring 












• 


• 






29 radiata (Aublet) Spring 


• 


• 


• 












• 


30 speeiosa A. Braun 












• 


• 






3 1 haenkeana Spring 












• 


• 


• 




32 hartwegiana Spring 












• 








33 viticulosa Klotzsch 




















34 pearcei Baker 














• 






35 meridens is Alston 




















36 popayanensis Hieron. 












• 




• 




37 cavifolia A. Braun 












• 








38 flacca Alston 




















39 macilenta Baker 












• 








40 mollis A. Braun 




















41 flagellata Spring 


• 


• 








• 


• 


• 


• 


42 moritziana Spring ex Klotzsch 












• 








43 cladorrhizans A. Braun 


• 
















• 


44 porphyrospora A. Braun 




















45 novae- hollandiae (Sw.) Spring 












• 




• 




46 chionoloma Alston ex Crabbe & Jermy 














• 


• 




47 lychnuchus Spring 





















238 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Table 1 (Continued) 







c 
.2 
'3 

O 

c 


B 

3 






u- 


C/} 


48 


ramosissima Baker 






49 


glossophylla Alston ex Crabbe & Jermy 






50 


minima Spring 


• 




51 


simplex Baker 






52 


tyleri A. C. Smith 






53 


cruciformis Alston ex Crabbe & Jermy 






54 


acanthostachys Baker 






55 


chrysoleuca Spring 






56 


contigua Baker 






57 


mendoncae Hieron. 






58 


leucoloma Alston ex Crabbe & Jermy 






59 


xiphophylla Baker 






60 


subs tip itata Spring 






61 


scalariformis A. C. Smith 






62 


wurdackii Alston 






63 


seemannii Baker 




• 


64 


erectifolia Spring 






65 


roraimensis Baker 






66 


scintillata Alston 






67 


flexuosa Spring 






68 


decomposita Spring 






69 


duidae A. C. Smith 






70 


euclimax Alston ex Crabbe & Jermy 






71 


vernicosa Baker 






72 


vestiens Baker 






73 


anisoclada Alston ex Crabbe & Jermy 






74 


microphylla (Kunth) Spring 






75 


schultesii Alston ex Crabbe & Jermy 






76 


microdonta A. C. Smith 






77 


jungermannioides (Gaudich.) Spring 






78 


arenaria Baker 






79 


brevifolia Baker 






80 


sandwithii Alston 




• 


81 


valdepilosa Baker subsp. valdepilosa 
subsp. tricholoma Jermy & Rankin 






82 


fragillima Silveira 






83 


ovifolia Baker subsp. philipsonii 
Jermy & Rankin 






84 


cabrerensis Hieron. 






85 


tuberculata Spruce ex Baker 




• 


86 


rhodostachya Baker 






87 


polar oensis Jen man 




• 


88 


muscosa Spring 






89 


dendncola Jen man 


• 


• 


90 


tenuissima Fee 






91 


falcata (Beauv.) Spring 


• 




92 


densifolia Spruce ex Hook. 






93 


kochii Hieron. 




• 


94 


truncata Karsten ex A. Braun 










3 ■§ 

N E 



3 U O 

o > u 



o 




a 






p 


> 


N 


3 






d 


o 


u 


o 


u 


UJ 


cu 


03 


03 






SELAGINELLA IN TROPICAL SOUTH AMERICA 239 

Table 1 (Continued) 







03 

c 
2 

'3 

O 

o 

c 
1) 


B 
a 
c 

3 


G 

3 


J2 

N 
C 

<u ■ 


.2 

JO 

B 

o 
o 


o 

a 
o 




.2 
'> 

"o 








Uu 


c/a 





> 


U 


U 


Oh 


CQ 


CQ 


95 


homaliae A. Braun 










• 








• 


96 


calosticha Spring 








• 






• 






97 


breynii Spring 










• 








• 


98 


applanata A. Braun 










• 




• 






99 


tarapotensis Baker 










• 


• 


• 


• 




100 


producta Baker 


• 


• 


• 


• 


• 




• 




• 


101 


macrostachya (Spring) Spring 


















• 


102 


revoluta Baker 


• 


• 


• 


• 


• 




• 




• 


103 


porelloides (Lam.) Spring 


• 




• 














104 


exaltata (Kunze) Spring 










• 


• 


• 




• 


105 


tomentosa Spring 










• 


• 








106 


conduplicata Spring 


• 


• 






• 




• 




• 


107 


asperula Spring 








• 


• 




• 


• 


• 


108 


fragilis A. Braun 








• 


• 








• 


109 


parkeri (Hook. & Grev.) Spring 


















• 


110 


arthritica Alston 










• 


• 








111 


geniculata (C. Presl) Spring 










• 


• 


• 






112 


dasyloma Alston 










• 


• 








113 


anaclasta Alston ex Crabbe & Jermy 








• 












114 


kunzeana A. Braun 








• 


• 


• 


• 






115 


lingulata Spring 










• 


• 


• 


• 




116 


articulata (Kunze) Spring 










• 


• 


• 






117 


valida Alston 




















118 


calceolata Jermy & Rankin 










• 










119 


marginata (Humb. & Bonpl.) Spring 








• 








• 




120 


atirrensis Hieron. 










• 




• 






121 


kraussiana (Kunze) A. Braun 




















122 


asplundii Alston ex Crabbe & Jermy 










• 




• 






123 


epirrhizos Spring 


• 


• 


• 














124 


poeppigiana (Spring) Spring ex Splitg. 












• 


• 






125 


suavis (Spring) Spring 














• 






126 


s ilves tr is Asplund 










• 


• 


• 


• 




127 


trisulcata Asplund 












• 


• 


• 




128 


sericea A. Braun 












• 








129 


sulcata (Desv.) Spring ex Martius 




• 












• 




130 


horizontalis (C. Presl) Spring 








• 


• 


• 








131 


microtus A. Braun 










• 


• 








132 


humboltiana A. Braun 








• 


• 










133 


diffusa (C. Presl) Spring 





















Leaf. In the upper branches of all species, leaves are seen as two kinds: median, on the upper 
(dorsal) plane of the 'frond', and lateral, on the lower (ventral) side. Those referred to as 
axillary leaves are modified lateral leaves and occur at all points of dichotomy. They are also 
of taxonomic value in regard to shape and margin characters. 

In dendroid species the leaves on the lower stem are spirally arranged and may not be 
readily designated medium or lateral. It is a general rule that the shape of both leaf types, but 



240 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

especially of the median leaves, is narrower near the branch apex, and the assessment of leaf 
shape should be made from those on primary branches. There is often considerable variation 
in density and length of cilia on leaves. Somers (1978) points out the importance of uni- or 
multicellular cilia in species delimitation. The degree of development of leaf auricles is 
variable. In species with two or more steles, the stems become sulcate on drying and auricles 
may be hidden in these furrows. The colour of leaves (either a deep or pale green) can be 
diagnostic. Blue iridescence, which is mainly an optical characteristic of the leaf epidermis, 
may be found in some species; but its development is usually affected by environmental 
conditions. The presence of white 'streaks' on leaves is not consistent and of no taxonomic 
use. The normally clearly defined opaque white borders to both median and lateral leaves 
are usually significant. The distribution of stomata and of papillae on leaf epidermises (the 
latter seen clearly in gum-chloral preparations) may prove to be useful diagnostic characters 
(see Mital, 1969; Bienfait & Waterkeyn, 1974) and needs further investigation. In a few 
species, the lateral leaves curl under the stems in a characteristic and diagnostic manner, 
but this feature may not be readily apparent if the specimen was moist when pressed. 

Spores. The colour of microspores has been elaborated upon by Hellwig (1969) but is not 
used here in detail. Somers (1978) found considerable variation within species, and that the 
colour of microspores may depend on the degree of maturation. These considerations also 
apply to the colour of megaspores. Sporoderm morphology of both micro- and megaspores 
is, on the other hand, diagnostic, and has been used by Hieronymus (1901), Hellwig (1969), 
and Somers (1978). Examination of spores with the scanning electron microscope may show 
micromorphological differences, but the significance of these can be over-emphasised and so 
obscure group similarities. However, sporoderm ornamentation can be diagnostic and may 
be used more usefully as an aid to identification than in this key which is constructed as far as 
possible for naming sterile material. 



Key to species 

la Leaves uniform, spirally arranged (subgenus Selaginella) 2 

1 b Leaves dimorphous, at least those on the ultimate branches, in 4 ranks 

(subgenus Stachygynadrum) 4 

2a(l) Leafy stems strongly dorsiventral, the zone of green leaves well developed on 
upper side of stem; leaves on all sides of stem with base decurrent or at least 
those on underside strongly decurrent; branch tips involute in dormant 
condition 3. peruviana (p. 251) 

2b Leafy stems radially symmetrical; leaves on all sides of stem equal in length, 

shape, position and texture, with bases abruptly adnate (rarely decurrent on 
main stem); branch tips not involute in dormant condition .... 3 

3a(2) Leaf-bases glabrous; leaf apex setae usually milk-white, opaque (if whitish to 

greenish-lutescentandsubtranslucent, then setae slightly attenuate) 1. sellowii (p. 250) 

3b Leaf-bases predominantly pubescent (if, rarely, mostly glabrous, then setae 
strongly attenuate); leaf apical setae whitish to greenish or lutescent, translucent 
tosubopaque 2. sartorii (p. 251) 

4a(l) Rhizophores ventral, i.e. arising on the same side as the axillary leaves; stems 

not articulate; mega- and microsporophylls the same size .... 5 

4b Rhizophores dorsal, i.e. arising on the opposite side of the stem to the axillary 
leaves; stems articulate; megasporophylls sometimes much larger than micro- 
sporophylls 129 (p. 248) 

5a(4) Stems tufted, curling inwards when dry: leaves rather thick and stitTin texture 6 

5b Stems not tufted, not curling when dry; leaves usually membranaceous . . 7 

6a(5) Median leaves not margined; texture subcoriaceous; old leaves dark brown 

4. convoluta (p. 251) 



SELAGINELLA IN TROPICAL SOUTH AMERICA 241 

6b Median leaves white-margined; texture not subcoriaceous; old leaves buff 

5. pallescens (p. 252) 

7a(5) Stems red or pink 8 

7b Stems stramineous or green 10 

8a(7) Lateral leaves minutely and evenly denticulate on upper margin 

8. haematodes (p. 254) 
8b Lateral leaves ciliate or ciliolate on upper margin (near base only) ... 9 

9a(8) Lateral leaves denticulate at apex on both sides, ciliolate at base on upper 
margin oblong-lanceolate; leaves usually dimorphic before first branch on main 
stem; median leaves with conspicuous white margin and arista half as long as 
lamina 6. erythropus (p. 253) 

9b Lateral leaves entire throughout the lower margin, long-ciliate at base on upper 
margin, ovate; leaves usually uniform throughout simple part of main stem; 
median leaves with narrow white margin, shortly acuminate-aristate 
7. umbrosa (p. 253) 

10a(7) Plants when fully developed with distinct, usually erect, main stem; 

leaves ± uniform, appressed at least in lower third 11 

10b Plants with no such distinct and erect stems but usually creeping; leaves for the 

most part entirely dimorphous throughout the plant 44 

lla(10) Main stem prostrate or arching 12 

lib Main stem erect 13 

1 2a( 11) Stems pubescent, terete; lateral leaves pale green, acuminate . 9. coarctata (p. 255) 
12b Stems glabrous, compressed; lateral leaves dark green, subobtuse 10. longissima (p. 255) 

1 3a( 11) Main stem compressed 14 

13b Main stem terete 15 

14a(13) Leaves on main stem normally uniform to internode above first branch; lateral 

leaves deltate to oblong, 4-6 mm long, herbaceous . 11. mazaruniensis (p. 255) 

14b Leaves on main stem dimorphous well below first branch; lateral leaves deltate, 

3 mm long, subcoriaceous 24. amazonica (p. 261) 

1 5a( 1 3) Leaves on simple part of main stem uniform nearly to first branch, and often 

to internode above 16 

15b Leaves on simple part of main stem uniform, but becoming dimorphous below 

the first branch 28 

16a(15) Lateral leaves on the main stem (in the middle of the frond), and on the oldest 

primary branches ovate-deltate, with a symmetrical apex .... 17 

16b Lateral leaves on main stem (in the middle of the frond), and on the oldest 

primary branches oblong, usually with an oblique apex 24 

1 7a( 1 6) Median leaves broadly elliptic-ovate, densely ciliate at least at base, apex blunt 

17. terezoana (p. 257) 
1 7b Median leaves variously shaped, not densely ciliate, apex acuminate to aristate 1 8 

1 8a( 1 7) Median leaves distinctly aristate 19 

18b Median leaves acuminate 21 

1 9a( 1 8) Leaves of main stem obliquely inserted, 3-6 mm long; median leaves long-aristate 

14. mortoniana (p. 256) 
19b Leaves of main stem above the first branch patent, up to 3 mm long; median 

leaves shortly aristate 20 

20a(19) Lateral branches mostly regularly pinnate, with simple branchlets; median 
leaves oblong-ovate, more or less equal-sided, with costa raised; lateral leaves 
regularly serrate 12. palmiformis (p. 256) 

20b Lateral branches mostly bipinnate; median leaves broadly elliptic, with broad 
outer side; with costa not raised; lateral leaves with few cilia at base, distantly 
serrate upwards 13. flabellata var. latifrons (p. 256) 



242 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

2 1 a( 1 8) Lateral leaves of branches more than 5 mm long 22 

21b Lateral leaves of branches up to 4 mm long 23 

22a(21) Branch lateral leaves 5x2 mm; branch system 2-or-more-pinnate; microspores 

with elongate papillae 15. lechleri (p. 256) 

22b Branch lateral leaves c. 6x2*5-3 mm; branch system 1-2-pinnate; microspores 

smooth 16. cheiromorpha (p. 257) 

23a(21) Lateral leaves glabrous beneath, c. 2 - 5 mm long .... 18. anceps (p. 257) 

23b Lateral leaves pubescent beneath, c. 4 mm long .... 19. hirsuta (p. 259) 

24a(16) Median leaves ± symmetrical, apex acuminate to aristate 25 

24b Median leaves hemi-orbicular, i.e. distictly asymmetrical, apex obtuse or acuminate 27 

25a(24) Branch system ovate or oblong in outline; lateral leaves distinctly dimidiate 

21. wolfii (p. 260) 
25b Branch system pedate in outline; lateral leaves not markedly dimidiate . . 26 

26a(25) Median leaves obovate-elliptic, dentate, apex acuminate often into a short arista 

20. bahiensis subsp. bahiensis (p. 259) 
26b Median leaves lanceolate-elliptic, denticulate, apex acute 

20. bahiensis subsp. manausensis (p. 260) 

27a(24) Median leaves sparingly ciliate at base, apex obtuse; lateral leaves 7-8 mm 

23. praestans (p. 260) 

27b Median leaves denticulate at base, apex acute; lateral leaves rarely more than 5 mm 

25. oaxacana (p. 261) 

28a(15) Largest lateral leaves 6-10 mm long 29 

28b Largest lateral leaves 3-4 mm long 35 

29a(28) Lateral leaves ciliate at base; lateral leaves of secondary and tertiary branches about 

equal 30 

29b Lateral leaves denticulate at base; lateral leaves of secondary and tertiary 

branches equal or unequal 33 

30a(29) Median leaves not aristate, at most slightly acuminate 31 

30b Median leaves aristate 32 

31a(30) Costa of median leaves not central; abaxial part of lamina twice as broad as 

adaxial part 23. praestans (p. 260) 

31b Costa of median leaves nearly central; abaxial part of lamina not twice as broad 

as adaxial part 26. quadrifaria (p. 261) 

32a(30) Median leaves longer than broad, ± asymmetrical, tapering gradually into a 

short arista 27. tanyclada (p. 263) 

32b Median leaves broader than long, strongly asymmetrical, abruptly acuminate 

with a ± long arista 28. bombycina (p. 264) 

33a(29) Lateral leaves of secondary and tertiary branches about the same size 

30. speciosa (p. 265) 
33b Lateral leaves of tertiary branches much smaller than those of secondary branches 34 

34a(33) Lateral leaves of upper part of main stem with conspicuous anterior basal lobe 
often overlapping stem; median leaves ± symmetrical, shortly aristate 

21. wolfii (p. 260) 

34b Lateral leaves of upper part of main stem without conspicuous anterior basal 

lobe; median leaves distinctly asymmetrical with long arista . 22. hartii (p. 260) 

35a(28) Lateral leaves of main stem subcoriaceous, imbricate, denticulate 

24. amazonica (p. 261) 
35b Lateral leaves of main stem herbaceous, imbricate or not, denticulate to ciliate 36 

36a(35) Lateral leaves ciliate at base 37 

36b Lateral leaves denticulate at base 41 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



243 



37a(35) 
37b 

38a(37) 
38b 

39a(38) 
39b 

40a(39) 

40b 

41a(36) 

41b 

42a(41) 

42b 
43a(42) 

43b 

44a(10) 

44b 

45a(44) 
45b 

46a(45) 
46b 



47a(46) 
47b 



48a(47) 

48b 
49a(48) 

49b 

50a(49) 

50b 

51a(50) 
51b 



Lateral leaves ovate with cilia evenly spaced, uniform; microspores papillose 

33. viticulosa (p. 
Lateral leaves ovate-oblong or lanceolate with cilia irregular in length and 
spacing; microspores papillose or not 

Median leaves of main stem distinctly asymmetrical, with an obtuse apex 
abruptly forming an arista ± as long as the leaf-blade . 31. haenkeana (p. 

Median leaves of main stem ± symmetrical, with an acute apex tapering to a 
arista <§ length of leaf-blade 

Median leaves with a short arista and large outer auricles; with no, or few, cilia 
at base; axillary leaves narrowly deltate or deltate-ovate; microspores smooth 

32. hartwegiana (p. 

Median leaves with an arista about half as long as leaf-blade, outer auricle 
virtually absent, ± densely ciliate at base; axillary leaves lanceolate; microspores 
papillate 

Lateral leaves 3-4 mm long, upper margin toothed with few short cilia at base 

35. meridensis (p. 
Lateral leaves 0*5-2 mm long, upper margins with long, often dense, cilia 

29. radiata (p. 

Median leaves aristate 

Median leaves not aristate 



35. meridensis (p. 



Branch system caulescent, main stem 15-2 mm thick; lateral leaves overlapping 

24. amazonica (p. 
Branch system diffuse, main stem not more than 1 mm thick; lateral leaves distant 

Median leaves broadly ovate to elliptic, obtuse with small acumen; lateral leaves 
oblong-elliptic 34. pearcei (p. 

Median leaves lanceolate-elliptic, acute or shortly acuminate; lateral leaves 
broadly ovate-elliptic 36. popayanensis (p. 

At least some stems and/or branches with flagelliform apices, or plants sobo- 

liferous 

Stems and/or branches without flagelliform apices; plants not soboliferous 

Mature stems rather stout, about 1 -5 mm in diameter . . 10. longissima (p. 
Mature stems slender, up to 1 mm in diameter 

Plant truly creeping, main stems bearing rhizophores above lower third . 
Plant tending to be erect or upper branches ascending, main stems usually 
without rhizophores above lower third 

Lateral leaves strongly ciliate, at least in lower half . . . 40. mollis (p. 
Lateral leaves not more than denticulate (occasionally with a few short cilia on 
auricles) 

Lateral leaves up to twice as long as broad; median leaves usually aristate 

41. flagellata (p. 
Lateral leaves not more than 1*5 times as long as broad; median leaves rarely aristate 

Median leaves cordate at base, nearly as broad as long, subacute; lateral leaves 
rounded at apex 76. microdonta (p. 

Median leaves not cordate at base, 15 times as long as broad, acute to acuminate 
or shortly aristate; lateral leaves acute or acuminate 

Median leaves acute or shortly and abruptly acuminate, entire or finely toothed, not 

papillate 37. cavifolia (p. 

Median leaves acuminate-aristate, often coarsely toothed, surface papillate 



Lateral leaves broadly ovate with an acuminate apex 
Lateral leaves lanceolate-elliptic, apex acute . 



38. flacca (p. 
39. macilenta (p. 



266) 
38 

265) 

39 

266) 

40 

267) 

264) 

267) 
42 

261) 
43 

267) 
269) 

45 

58 

255) 
46 

47 

52 
272) 

48 



272) 
49 



288) 
50 



269) 
51 

269) 

272) 



52a(46) Lateral leaves acuminate to aristate; median leaves ciliate 54. acanthostachys (p. 278) 



244 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

52b Lateral leaves not aristate; median leaves not ciliate, although with occasional 

long teeth in upper part 53 

53a(52) Median leaves not auriculate; lateral branches with flagelliform apices; basal 

soboles not present 54 

53b Median leaves auriculate; lateral branches rarely with flagelliform apices 

{acanthostachys); basal soboles usually present 55 

54a(53) Lateral leaves ovate, usually with short hairs on basiscopic side of upper surface 

and with two false nerves; megaspores orange . . 44. porphyrospora (p. 274) 

54b Lateral leaves linear-lanceolate or oblong, glabrous and with a single nerve; 

megaspores pale cream 43. cladorrhizans (p. 274) 

55a(53) Median leaves not bordered, finely denticulate . 42. moritziana (p. 273) 

55b Median leaves distinctly white- or hyaline-bordered, dentate to often spinose- 

toothed or ciliate 56 

56a(55) Leaves spinose-dentate but not ciliate, lateral leaves with conspicuous white 
border often 2-3 cells broad on acroscopic margin, drying ± flat 
46. chionoloma (p. 276) 

56b Leaves ciliate on acroscopic margin, with narrow ± hyaline border, curling 

when dry 57 

57a(56) Upper lateral leaves acuminate to aristate; median leaves with long white arista 

54. acanthostachys (p. 278) 
57b Upper lateral leaves at the most only apiculate; median leaves without white arista 

45. novae-hollandiae (p. 274) 

58a(44) Main stems usually suberect without rhizophores above lower third ... 59 

58b Main stems prostrate with rhizophores above the lower third .... 94 

59(58) Sporophylls strongly dimorphous 60 

59b Sporophylls uniform or nearly so 65 

60a(59) Lateral leaves obtuse, oblong 61 

60b Lateral leaves acute, variable in shape 63 

61a(60) Lateral leaves ciliolate or ciliate at base 47. lychnuchus (p. 276) 

61b Lateral leaves denticulate at base 62 

62a(61) Stem 1 mm in diameter at base; lateral leaves broadest below middle, more than 

twice as long as broad 48. ramosissima (p. 277) 

62b Stem filiform, i.e. less than 1 mm diameter throughout; lateral leaves broadest 

above middle, up to twice as long as broad .... 49. glossophylla (p. 277) 

63a(60) Median leaves long-aristate; lateral leaves with distinct white border 

46. chionoloma (p. 276) 
63b Median leaves acute to acuminate; lateral leaves not bordered .... 64 

64a(63) Lateral leaves oblong, ciliate; lower sporophylls ciliate . . 50. minima (p. 277) 
64b Lateral leaves ovate, denticulate; lower sporophylls denticulate . 51. simplex (p. 277) 

65a(59) Lateral leaves densely pubescent on both surfaces or shortly so on upper surface 

of lower half 66 

65b Leaves glabrous on both surfaces 67 

66a(65) Both median and lateral leaves with long hairs on surface and on margins as 

cilia; lateral leaves with single nerve 52. tyleri (p. 277) 

66b Median leaves glabrous, lateral leaves with short hairs on basiscopic side of 

upper surface only; lateral leaves with two false nerves 44. porphyrospora (p. 274) 

67a(65) Stems 10-15 mm across (including leaves) 68 

67b Stems less than 8 mm across (including leaves) 71 

68a(67) Median leaves subacute 53. cruciformis (p. 278) 

68b Median leaves acuminate to aristate 69 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



245 



69a(68) 
69b 

70a(69) 
70b 

71a(67) 
71b 

72a(71) 

72b 

73a(72) 
73b 

74a(71) 
74b 

75a 
75b 

76a 
76b 

77a 
77b 

78a(77) 
78b 

79a(78) 
79b 

80a(79) 
80b 

81a(80) 

81b 

82a(79) 
82b 

83a(82) 
83b 

84a(82) 
84b 

85a(84) 

85b 
86a(85) 

86b 



Median leaves finely toothed, without cilia 
Median leaves ciliate 

Median leaves with a distinct arista . 
Median leaves acuminate 

Median leaves ciliate at least in the lower half 

Median leaves not ciliate (although sometimes sharply toothed) 



55. chrysoleuca (p. 

56. contigua (p. 
57. mendoncae (p. 



278) 
70 

278) 
279) 

72 
74 



Sporophylls acuminate, ciliate throughout margin; leaves dark green; lateral 
leaves ovate, symmetrical 54. acanthostachys (p. 278) 

Sporophylls subacute, toothed, sometimes ciliate at base; leaves light green; 
lateral leaves asymmetrical 73 



Median leaves subacute .... 
Median leaves aristate or abruptly acuminate 



58. leucoloma (p. 279) 
45. novae-hollandiae (p. 274) 



Lateral leaves oblong-lanceolate, more than twice as long as broad, apex obtuse; 

stems not filiform, usually more than 0'5 mm in diameter .... 75 

Lateral leaves ovate deltate to broadly lanceolate, up to 15 times as long as 

broad, apex acute; stems filiform, up to 0*25 mm in diameter (except S. pearcei) 87 

Arista of median leaves more than half as long as lamina 76 

Arista of median leaves up to a third as long as lamina, or wanting, or leaves 
acuminate 77 



Lateral leaves closely and regularly spaced, dark green 
Lateral leaves distant, light green .... 

Lateral leaves distinctly white-margined . 

Lateral leaves not white-margined .... 



59. xiphophylla (p. 

60. substipitata (p. 

84. cabrerenis (p. 



Lateral leaves transversely disposed in opposite plane to stem; median leaves 
elliptic-orbicular, closely ciliolate . . . . 61. scalariformis (p. 

Lateral leaves not as above; median leaves oblong-ovate to obovate or narrowly 
elliptical, dentate 

Sporophylls lanceolate-deltate 

Sporophylls ovate-deltate 



279) 
279) 

294) 

78 

280) 

79 

80 
82 



62. wurdackii (p. 280) 



Stems slender, 0*5 mm in diameter; lateral leaves ± entire . 

Stems stout, 1 mm in diameter; lateral leaves ciliate or toothed .... 

Median leaves oblong-lanceolate, acuminate with short hyaline tip; lateral 
leaves 2-3 mm long, patent even when dry .... 63. seemannii (p. 

Median leaves ovate-elliptic, acuminate into arista; lateral leaves 1-2 mm long, 
especially the lower ones erect and clasping the stem . 64. erectifolia (p. 

Median leaves acute 

Median leaves acuminate or aristate 



66. scintillata (p. 
65. roraimensis (p. 



Lateral leaves c . 1 mm long; rhizophores short and filiform . 
Lateral leaves c. 2 mm long; rhizophores long and stiff . 

Main stem (incl. lateral leaves) 8-10 mm wide; lateral leaves entire 67. flexuosa (p. 
Main stem (incl. lateral leaves) <5mm wide, or if wider then lateral leaves 
wide-spaced; lateral leaves denticulate at least on acroscopic edge . 

Lateral leaves on main branches distant, with hyaline teeth on both margins 

68. decomposita (p. 
Lateral leaves on main branches closely set, with teeth only on acroscopic margin 

Median leaves abruptly acuminate into short, ± hyaline arista; lateral leaves 

dentate towards acute apex 69. duidae (p. 

Median leaves acuminate; lateral leaves with distinct teeth only at base, apex obtuse 

70. euclimax (p. 



87a(74) Surface of leaves polished and texture firm 



280) 

282) 

83 
84 

282) 
282) 

282) 
85 

285) 
86 

286) 

286) 
88 



246 

87b 

88a(87) 
88b 

89a(88) 
89b 

90a(89) 
90b 

91a(87) 

91b 

92a(91) 

92b 

93a(92) 
93b 

94a(58) 
94b 

95a(94) 
95b 

96a(95) 
96b 



97a(96) 
97b 

98a(97) 
98b 

99a(98) 
99b 

100a(99) 
100b 

101a(97) 
101b 

102a(101) 
102b 

103a(101) 
103b 



A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Surface of leaves not polished and/or texture somewhat diaphanous . 

Lateral leaves all overlapping; lateral branches usually ascending 71. vernicosa (p. 
Lateral leaves spaced towards base of stem 

Lateral branches ascending 66. scintillata (p. 

Lateral branches usually spreading at an angle of 45 degrees .... 

Lateral leaves ovate, ± entire; median leaves subacute 86. rhodostachya (p. 

Lateral leaves broadly lanceolate, ciliatodentate; median leaves acuminate 

72. vestiens (p. 

Lateral leaves of main stem very distant and much longer than those of branches 

73. anisoclada (p. 
Lateral leaves of main stem less than one leafs length apart, not much longer 
than those of largest branches 

Lateral leaves ovate-deltate, distinctly asymmetrical . . 80. sandu ithii (p. 
Lateral leaves ovate-lanceolate to elliptic, ± symmetrical 

Median leaves lanceolate-elliptic, acute or shortly acuminate; lateral leaves 
broadly ovate-elliptic 36. popayanensis (p. 

Median leaves broadly ovate-elliptic, obtuse but with acute acumen; lateral 
leaves oblong-elliptic 34. pearcei (p. 

Lateral and axillary leaves ovate, up to twice as long as broad .... 
Lateral and axillary leaves oblong, usually more than twice as long as broad . 

Lateral leaves ciliate at base 

Lateral leaves denticulate 



Lateral leaves inserted obliquely, clasping the stem when dry; median leaves 
similar to lateral leaves suborbicular, subacute ... 74. microphylla (p. 

Lateral leaves patent, often rolling under from apex to base when dry; median 
leaves different from the lateral leaves broadly elliptic to ovate-lanceolate, 
subacute to apiculate 

Median leaves aristate, acuminate or cuspidate; lateral leaves acute or obtuse . 
Median leaves not aristate or acuminate or cuspidate; lateral leaves obtuse 

Median leaves with long aristate tip; lateral leaves c. 1 mm long 75. schultesii (p. 
Median leaves acuminate or cuspidate; lateral leaves 1 5—30 mm long 

Width of stem plus lateral leaves 3^ mm; secondary branches unbranched; 

rhizophores throughout main stem ... 77. jungermannioides (p. 

Width of stem plus lateral leaves 2-0-2-5 mm; secondary branches mostly 

dichotomous; rhizophores in lower third of main stem only .... 

Lateral leaves often densely ciliate on both margins, at least below; median 
leaves broadly elliptic, not acuminate 78. arenaria (p. 

Lateral leaves sparsely ciliate on acroscopic margin, basiscopic margin ± entire; 
median leaves ovate-elliptic, acuminate .... 79. brevifolia (p. 

Median leaves suborbicular, ciliate 

Median leaves lanceolate or oblong-elliptic, not ciliate (or occasionally so at apex) 

Lateral leaves 1 -5 times as long as broad, contiguous 

8 1 . valdepilosa subsp. valdepilosa (p. 
Lateral leaves twice as long as broad, not contiguous 

81. valdepilosa subsp. tricholoma (p. 

Median leaves lanceolate with distinct white border, without apical setae 

82. fragillima (p. 
Median leaves oblong-elliptic without white border, with 2-3 apical setae 

83. ovifolia subsp. philipsonii (p. 



91 

286) 
89 

282) 
90 

296) 
286) 

287) 

92 

289) 
93 

269) 

267) 

95 
114 

96 
104 

287) 



97 

98 
101 

288) 
99 



288) 
100 

289) 

289) 

102 
103 

292) 
292) 

294) 
294) 



1 04a(95) Median leaves subacute, neither aristate or acuminate 



105 



104b 
105a(104) 

105b 

106a(105) 
106b 

107a(106) 
107b 

108a(104) 
108b 

109a(108) 
109b 

110a(109) 
110b 

llla(HO) 
111b 

112a(109) 

112b 

1 13a(l 12) 

113b 

114a(94) 
114b 

1 15a(l 14) 

115b 

1 16a(l 15) 

116b 



SELAGINELLA IN TROPICAL SOUTH AMERICA 
Median leaves acuminate or anstate 



Upper surface of lateral leaves beset with tubercles near the lower margin 

85. tuberculata (p. 
Upper surface of lateral leaves smooth 



86. rhodostachya (p. 



Lateral leaves costate to apex .... 
Lateral leaves ecostate, or costate at base only 

Median leaves deltate-orbicular 

Median leaves ovate-oblong .... 

Median leaves (exclusive of the apical cusp) 

leaves, outer side cordate .... 
Median leaves (exclusive of the apical cusp) more than half as long as lateral 

leaves, outer side rounded or auricled . 



ess than 



76. microdonta (p. 
37. cavifolia (p. 

half as long as lateral 
87. potaroensis (p. 



Sporophylls uniform or nearly so; median leaves aristate 
Sporophylls dimorphous; median leaves acute to shortly aristate 

Lateral leaves ovate to ovato-deltate, 2- 5-3 mm long, apex becoming acute 

88. muscosa (p. 
Lateral leaves of main stem ovato-suborbicular, c. 1 mm long, apex rounded 
obtuse (those on fertile branches may be ovate-elliptic and < 2 mm long) 

Lateral leaves heteromorphous (i.e. some larger and closer than others); median 

leaves ovate-elliptic, apex acute 89. dendricola (p. 

Lateral leaves not heteromorphous; median leaves suborbicular-elliptic, apex 
aristate 

90. tenuissima (p. 

Lateral leaves obtuse or subacute, strongly cordate at base on upper side only 

103. porelloides (p. 
Lateral leaves acute or acuminate, rounded at base 



37. cavifolia (p. 
38. flacca (p. 



Sporophylls ovate; median leaves acuminate .... 
Sporophylls lanceolate; median leaves shortly aristate . 

Lateral leaves with hyaline margin of three or more rows of elongate cells on both 
sides of the leaf 

Lateral leaves with no hyaline margin; marginal cells on lower side of leaf near 
apex quadrate, or, if elongate, then cells with chloroplasts .... 

Median leaves ovate or ovate-deltate, acute-acuminate, falcate .... 
Median leaves oblong-lanceolate, aristate, not falcate 

Median leaves acuminate, ciliate for 2/3 of margin, apices pointing outwards; 

lateral leaves ciliate at base 91. falcata (p. 

Median leaves acute, ciliate on auricles only, apices pointing inwards; lateral 

leaves with at most a few cilia at base 92. densifolia (p. 

1 1 7a( 115) Stems (including leaves) up to 3*5 mm across, ascending, with short often slender 
rhizophores; lateral leaves densely ciliate, ensheathing stem when dry 

84. cabrerensis (p. 

117b Stems (including leaves) normally 6 mm or more across, prostrate, with long 
stout rhizophores; lateral leaves spinose-dentate, becoming ciliolate on auricles, 
not curling round stem when dry 93. kochii (p. 

1 1 8a( 114) Median leaves subacute or at most slightly cuspidate 

118b Median leaves aristate 

1 1 9a( 1 1 8) Stems (including leaves) 6-8 mm across; leaves sparsely and minutely toothed 

94. truncata (p. 
1 1 9b Stems (including leaves) up to 3*5 mm across; leaves spinose-toothed 

95. homaliae (p. 

1 20a( 1 1 8) Median leaves conspicuously white-margined 



247 
108 



296) 
106 

296) 
107 

288) 
269) 



297) 

109 

110 
112 



297) 
111 

299) 

299) 

305) 
113 

269) 
269) 

115 

118 

116 

117 

300) 
300) 

294) 

301) 

119 
120 

301) 

302) 
121 



248 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

120b Median leaves not conspicuously white-margined 124 

12 la(l 20) Lateral leaves not ciliate 122 

121b Lateral leaves ciliate 123 

1 22a( 121) Median leaves ovate-lanceolate, gradually attenuate into a long arista which 

equals lamina 59. xiphophylla (p. 279) 

122b Median leaves broadly oblong, abruptly attenuate, arista shorter than lamina 

96. calosticha (p. 302) 

1 23a( 121) Stems (including leaves) 7 mm or more across; microspores with capitate verrucae 

97. breynii (p. 302) 
1 23b Stems (including leaves) up to 5 mm across; microspores with elongate papillae 

98. applanata (p. 303) 

124a(120) Microspores with stalked papillae; arista of median leaves ± equalling lamina 125 

124b Microspores smooth; arista of median leaves not more than half lamina . . 126 

125a(l 24) Lateral leaves ovate-elliptic, twice as long as broad . . 99. tarapotensis (p. 303) 

125b Lateral leaves oblong-elliptic, about three times as long as broad 67. flexuosa (p. 282) 

1 26a( 1 24) Lateral leaves denticulate, cilia if present confined to auricles, subobtuse, drying flat 127 

126b Lateral leaves ciliate, subacute, often longitudinally inrolled .... 128 

1 27a(l 26) Branches usually short and simple; lateral leaves 2-3 mm long, not uniform in 
length and more distant in some parts of the stem; leaf texture membranous 

89. dendricola (p. 299) 

127b Branches usually pinnate; lateral leaves 3-4 mm long, uniform in both length 

and distribution; leaf texture firm 100. producta (p. 303) 

128a(126) Secondary branch units 4-10 cm, 3-4 times forked . 101. macrostachya (p. 304) 

128b Secondary branch units 1-2 cm, 1-2 times forked . . . 102. revoluta (p. 304) 

129a(4) Branches pubescent (main stems sometimes glabrescent) 130 

129b Branches glabrous 132 

130a( 1 29) Leaves of main stem spreading, rather close, obtuse . . 1 16. articulata (p. 313) 

130b Leaves of main stem directed upwards, distant, acute 131 

1 3 la( 1 30) Stem scandent with spreading, bipinnate, frond-like branches 104. exaltata (p. 306) 

131b Stem erect with an apical 3-4-pinnate frond-like portion 105. tomentosa (p. 306) 

1 32a( 1 29) Main stems simple, erect, with leaves subequal towards the base; lateral leaves 

appressed to stem 133 

132b Main stems branched from the base, prostrate or rarely ascending; with leaves 

dimorphous throughout; lateral leaves spreading 138 

1 33a( 1 32) Axillary leaves with long spur-like auricles . . . 106. conduplicata (p. 306) 

133b Axillary leaves exauriculate or with very short auricles 134 

1 34a( 1 33) Main stems with one stele 135 

134b Main stems with two steles 137 

1 35a( 1 34) Primary branch width (incl. leaves) 3-4 mm .... 107. asperula (p. 307) 

135b Primary branch width (incl. leaves) 5-7 mm 136 

1 36a( 1 35) All aerial stems determinate, not rooting at apex . . . 109. parkeri (p. 309) 
1 36b Main and primary branch stems indeterminate, usually rooting at apex 

108. fragilis (p. 308) 

1 37a( 1 34) Lateral leaves up to 2 mm broad, with a triangular auricle on lower side; outer 

auricles on median leaves usually rounded . . . . 111. geniculata (p. 310) 

137b Lateral leaves up to 3*5 mm broad, rounded or truncate on lower side at base; 

outer auricle on median leaves usually distinctly angular . 1 10. arthritica (p. 310) 

1 38a( 1 32) Median leaves obtuse, fringed with spreading hairs . 1 12. dasyloma (p. 311) 

138b Median leaves acute, acuminate or aristate, not fringed with spreading hairs 139 



SELAGINELLA IN TROPICAL SOUTH AMERICA 249 

1 39a( 1 38) Median leaves aristate, fringed with stiff spreading teeth, tipped by cilia; 

branches recurved 113. anaclasta (p. 312) 

139b Median leaves variable but with cilia at base only or wanting; branches not 

recurved 140 

1 40a( 139) Axillary leaves not conspicuously auriculate or ciliate, but sometimes peltate . 141 

140b Axillary leaves conspicuously auriculate or ciliate, or both 153 

14 la(140) Lateral leaves tapering from base to apex, with a conspicuous auricle on lower 

margin 114. kunzeana (p. 312) 

141b Lateral leaves parallel-sided or broadest in the middle, without a conspicuous 
lower auricle, sometimes truncate at base, or with an auricle on the upper side 
only 142 

1 42a( 141) Median leaves with two equally long auricles, the outer broad and the inner narrow 1 43 
142b Median leaves peltate or mono- or bi-auriculate (if latter then auricles of equal 

size and shape) 144 

143a(142) Median leaves acute 115. lingulata (p. 312) 

143b Median leaves long-aristate 117. valida (p. 313) 

144a(142) Median leaves acute 145 

144b Median leaves acuminate, cuspidate or aristate 148 

145a(144) Median leaves membranous 118. calceolata (p. 314) 

145b Median leaves chartaceus 146 

1 46a( 145) Median leaves peltate or with two long broad auricles; stems with one stele 

119. marginata (p. 314) 
146b Median leaves with one large outer auricle; stems with two steles ... 147 

147a(146) Lateral leaves 1 5-2-0 ( — 3) mm; median leaves with two auricles 120. atirrensis (p. 317) 
147b Lateral leaves 2-5-3 mm; median leaves with outer auricle only 121. kraussiana (p. 317) 

148a(144) Ultimate segments of branch system unequally dichotomous or pinnate; axillary 

leaves elliptic 149 

148b Ultimate segments of branch system equally dichotomous, flabellately arranged; 

axillary leaves not elliptic 151 

149a(148) Median leaves aristate, distinctly coarsely toothed; lateral leaves ovate-elliptic, 

toothed on the acroscopic margin 122. asplundii (p. 317) 

149b Median leaves acuminate or cuspidate, entire to denticulate; lateral leaves 

oblong-elliptic, entire to denticulate 150 

1 50a( 1 49) Median leaves acuminate with equal auricles; base of lateral leaves cordate 

123. epirrhizos (p. 318) 
1 50b Median leaves long-acuminate or cuspidate with unequal auricles; base of lateral 

leaves with narrow ciliate auricle on acroscopic side . 124. poeppigiana (p. 318) 

151a(148) Lateral leaves symmetrical and oblong in outline with rounded or very obtuse 

apex; axillary leaves narrowly oblanceolate ... 126. silvestris (p. 319) 

151b Lateral leaves asymmetrical in outline, broadest nearer the base, with pointed 

apex; axillary leaves ovate or lanceolate 152 

1 52a( 151) Lateral leaves distinctly toothed at least on acroscopic margin, ± symmetrical, 

midrib ± median; median leaves unbordered 127. trisulcata (p. 320) 

152b Lateral leaves indistinctly toothed, distinctly asymmetrical, midrib closer to 

posterior margin; median leaves with white borders . . 125. suavis (p. 319) 

153a(140) Main stems flagelliform at apex 129. sulcata (p. 321) 

153b Main stems not flagelliform at apex 154 

154a(153) Stems up to 15 mm across (including leaves), dichotomously branched and 
flabellate at apex; axillary leaves with two incurved entire auricles; median 
leaves often paler than lateral leaves 155 

154b Stems up to 8(-12)mm across (including leaves), unequally dichotomous or 



250 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

pinnately branched; axillary leaves with straight or curved dentate to ciliate 
auricles; median leaves same colour as lateral leaves 156 

1 55a(l 54) Lateral leaves with two auricles 128. sericea (p. 321) 

155b Lateral leaves with one auricle 116. articulata (p. 313) 

1 56a( 1 54) Lateral leaves narrowly oblong; median leaves aristate with one peltate auricle 1 5 7 

156b Lateral leaves elliptic, elliptic-lanceolate or deltate-ovate; median leaves acute or 

acuminate, usually with two auricles (the inner one sometimes rudimentary) 1 60 

1 57a( 1 56) Axillary leaves with long straight subparallel auricles curved at the tips 

129. sulcata (p. 321) 
157b Axillary leaves with short usually wholly curved auricles 158 

1 58a(l 57) Auricles on upper side of lateral leaves longer than those on the lower side, with 

cilia 1 -celled 130. horizontalis (p. 322) 

158b Auricles of lateral leaves subequal, with cilia one or several-celled ... 159 

1 59a( 158) Lateral leaves obtuse, distant, 3^ leaf-breadths apart on main stem 

131. microtus (p. 323) 
1 59b Lateral leaves subacute, close, 1-2 leaf-breadths apart on main stem 

132. humboldtiana (p. 323) 

160a(l 56) Lateral leaves c.\ '5 mm, lacking posterior auricle . . 120. atirrensis (p. 317) 

1 60b Lateral leaves (2-) 3 -5-5 mm, with ± triangular posterior auricle (often appressed to 

dorsal side of stem) 133. diffusa (p. 323) 



Species 
Subgenus SELAGINELLA 

1. Selaginella sellowii Hieron. in Hedwigia 39 : 306 (1900); Tryon in Ann. Mo. hot. Gdn 42 : 
34 (1955). Type from Brazil, Praia de San Diego, anno 1821, Sellow (B, fragment; NY). 

Selaginella rupestris f. brasiliensis Milde, Fil. Eur. Atlant. : 263 (1867), p.p. Type as above. 

Selaginella rupestris f. amazonica Milde, Fil. Eur. Atlant. : 263 (1867). Type from Peru. 

Selaginella rupestris sensu Sodiro, Crypt. Vase. Quit. : 592 (1893), non (L.) Spring (1838). 

Selaginella amazonica (Milde) Hieron. in Hedwigia 39:310 (1900), non S. amazonica 
Spring ( 1 840). Type from Peru. 

Selaginella mildei Hieron. in Engl. & Prantl, Nat. Pflanzenf. 1 (4) : 671 (1902). Type from 
Peru. 

Specimens seen: 

VENEZUELA. Lara: near Barquisimeto, 500 m, Alston 6382 (BM). 

COLOMBIA. Narino: Patia Valley, 700— 1200 m, Lehmann KK97 (K US); Rio Guaitara, Pasto, 

1 500 m, Lehmann BT776 (K, US). 

ECUADOR. Without exact locality: Sodiro s.n. (B, NY); Valle del Juanambu, Andre 2891 (NY); 

Quitonian Andes. Jameson 12 & 13 (BM); 'Valleys of the Andes', 1800-2100 m, Jameson 426 (BM, 

US). Imbabura: between Chota and Ambuqui, Fagerling & Wibom 1333 (S). Pichincha: near Perucho, 

1700 m.Asplund 20307 (S). 

PERU. Amazonas/Loreto: banks of Rio Maranon, near Tomependa, Bonpland s.n. (B, holotype; BM, 

isotype of 5. rupestris f. amazonica Milde). Cajamarca: Jaen, Penco, Raimondi 652 (BM). 

BOLIVIA. Cochabamba: Cochabamba, 2600 m, Tutin 1607 (BM); Cerveceria Colon, 2700 m, Balls 

B. 6229 (US); gravelly, sparsely shrub-covered slope, 2520 m, 3 miles W Quillacello-Cochabamba, 

Hermann 24595 (BM). La Paz: Neuvo mundo, Troll 898 (B); Muru Muntani, Larecaja, 

\200 m, Williams 1400 (NY, US). Santa Cruz: Resinas del Fuerte, Samaipata, 2000 m, Steinbach 

8254 (NY). Tarija: Toldos, 3200 m, Fiebrig 3322 (BM). 

BRAZIL. Without exact locality: type as above. Bahia: near Machado Portello, Rose & Russell 19937 

(NY). Rio de Janeiro: Corcovado, v. Lutzelburg 18870 (NY); Lindman, A 213 (BM); Nictheroy 

Frubuky, Brade 12825 (BM); above garden of British Embassy, Rio Cosme valles 95, Lutz 1883 (BM). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 251 

Rio Grande do Sul: Santa Angelo, Jurgens L 30 (BM); Bohmeback, Jurgens L 29 (BM, NY); 
Montenegro, Dutra 1370 (BM). 

Geographical range: Cuba and Mexico to Argentina. 

Notes: Schomburgk (Reisen Brit. Guiana 3:794, 880 (1948)) reports S. rupestris from 
British Guiana: 'An den Ufern des Demerara auf lichten Waldstellen'. No one has found 
any species of this group in the Guianas since, but the record may refer to S. sellowii or S. 
s art or ii. 

2. Selaginella sartorii Hieron. in Hedwigia 39 : 304 (1900); Knox in Trans, hot. Soc. Edinb. 
35 : 245 (1950); Tryon in Ann. Mo. hot. Gdn 42 : 36 (1955). Type from Mexico, Sartorius 
s.n.(B). 

Seleginella rupestris f. brasiliensis Milde, Fil. Eur. Atlant. : 263 (1867), p.p. Type from 

Venezuela. 
Seleginella sartorii var. venezuelensis Hieron. in Hedwigia 39:305 (1900). Type from 

Venezuela. 
Seleginella rupestris sensu Kunth in Reprium Spec. nov. Reg. Veg. 43 : 94 (1926), non. (L.) 

Spring. 

Specimens seen: 

VENEZUELA, Merida: Langunillas, Marcuzzi s.n. (BM); near Ejido, 1700 m, Alston 6114 (BM); 

Paramo de Mucuti, Mucuchies, Moritz 370 (BM, K, isotypes of S. sartorii var. venezuelensis Hieron.). 

Tachira: near la Grita, 1100 m, Alston 7051 (BM). Trujillo: between Valera and Timotes, 1800 m, 

Alston 6507 (BM), 65 1 3 (BM). 

COLOMBIA. Cundinamarca: Soacha, 2600 m, Lindig 1523 (BM). Norte de Santander: Paramo de 

Fontibon, Pamplona, 2500 m, Alston 7285a (BM). Santander: Malaga, 2200 m, Alston 7390 (BM). 

BOLIVIA. La Paz: Sorata, 2400 m, Rusby 462a (NY). 

Geographical range: Mexico to Colombia, possibly to Bolivia. 

Note: The specimen from Bolivia (Rusby 462a) probably belongs here, and we would like to 
see more material from this area. 

3. Selaginella peruviana (Milde) Hieron. in Hedwigia 39 : 307 (1900); Knox in Trans, bot. 
Soc. Edinb. 35 : 244 (1950); Tryon in Ann. Mo. bot. Gdn 42 : 77 (1955). Type from Peru, 
Huanuco, Ruiz s.n. (NY). 

Selaginella rupestris f. peruviana Milde, Fil. Eur. Atlant. : 263 (1 867). Type as above. 
Lycopodioides rupestris sensu Kuntze, Rev. Gen. 3 : 379 (1 89 1 ), non (L.) Kuntze. 

Specimens seen: 

ECUADOR. Without precise location: Nabon, Rose 23039 (NY). Loja: bank of Rio Arenal, Puente 

Buqueron, S W of Catamayo, Asplund 1 8064 (S). 

PERU. Without precise locality, Diehl25\l (BM), 2565 (BM). Cuzco: Urubamba Valley, 150 m, Tutin 

1359 (BM). Huanuco: type as above. Junin: Orora, Rose 19468 (NY). Lima: Matucana, 2400 m, 

Wilkes s.n. (NY); 2400 m, Asplund 1 1093 (S). 

BOLIVIA. Without precise locality: Bang 1 1 1 (NY); Kuntze s.n. (NY). La Paz: Sorata, Mandon 1531 

(BM); vicinity La Paz, 3000-3450 m, Cardenas 46 (NY); Rose 18845 (NY); Williams 2649 (NY); Bang 

1 20 (NY); Obrajas near La Paz, 3300 m, Buchtien 4299 (NY); Talca Chugiaguillo, Bang 787 p.p. (NY). 

Geographical range: Oklahoma and New Mexico, south to Argentina. 



Subgenus STACHYGYNANDRUM 

4. Selaginella convoluta (Arnott) Spring in Martius, Fl. Bras. 1 (2) : 131 (1840), in Mem. 
Acad. r. Sci. Lett. Belg. 24 : 69 (1850); R. Schomburgk, Reise, Brit. Guiana 3 : 880 (1848); 
Baker, Fern Allies : 88 ( 1 887); Maury in J. Bot., Paris 3 : 1 30 ( 1 889). Type from Brazil, Rio 
de Janeiro, Jameson s.n. (E). 



252 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Lycopodium convolutum Arnott in Mem. Wernerian nat. Hist. Soc. 5:199 (1824); 

reimpr. in Mem. Soc. Hist. nat. Paris 1 (2) : 327 (1 824). Type as above. 
Lycopodium convolutum (Beauv.) Desv. (1814), nom. illeg. [published only in synonymy]. 
Lycopodium bryoptehs sensu Aublet, Hist. PL Guiane Franc. 2 : 967 (1775), non L. 
Lycopodium hygrometricum Martius, Reise in Bras. 2 : 792 (1828). Type from Brazil. 
Lycopodium revolutum Hook & Grev. in Hook., Bot. Misc. 2:381 [ 4 28 1 '] (1 83 1 ). 
Selaginella hygrometrica (Martius) Spring ex Martius in Flora, Jena (Beibl.) 1837 (2) : 126 

(1837)/Type as above. 
Selaginella orbignvana Spring ex Decaisne, Arch. Mus. 2 : 194 (1 84 1^42), nom. nud.; in 

Bull. Acad. r. Belg. 10 : 136 (1843); in Mem. Acad. r. Sci. Lett. Belg. 24 : 68 (1850). Type 

from Bolivia. 
Selaginella longispicata Underw. in Pubis Field Mus. (Bot.) 1 : 287 (1896); Hieron. in 

Hedwigia 63 : 286 (1917). Type from Mexico. 

Specimens seen: 

GUYANA. Demerara: without further locality, Hancock s.n. (K). 

VENEZUELA. Without precise locality: Stevens s.n. (NY). Lara: near Barquisimeto, 500 m, Alston 

6383 (BM). 

COLOMBIA. Magdalena: Santa Marta, Pur die s.n. (BM); Valle Dupar, Haught 2338 (BM, COL); San 

Sebastian, Sierra Nevada de Santa Marta, 6200 m, Foster & Smith 1512 (COL). 

BOLIVIA. Santa Cruz: Yuanatos, Prov. Chiquitos, D'Orbigny 263 (P, type of S. orbignyana). 

BRAZIL. Without exact locality: Redondo s.n. (BM); Glaziou 1217 (C). Bahia: near Machado Portello, 

Rose & Russell 20012 (NY); Barlima, Rose & Russell 19796 (NY); Caetano-Nazareth, Pernambuco, 

Werdermann 2826 (B). Minas Gerais: near Juiz de Fora, Damazio 727 (P). Paraiba: Serra d'Olho 

D'Agua, Buquarao, Luetzelburg 287 (M). Piaui: Catingas Sertao, Gardner 2020 (BM). Rio de Janeiro: 

Regnell 368 (C); Glaziou s.n. (C); Serra Estrella, Frade, Luetzelburg 272 (B, M); Leme, Luetzelburg 52 

(M). Sao Paulo: Usteri 26a (P); Salto Grande do Paranapanema, Edwall 4950 (S). 

Geographical range: Guatemala and Greater Antilles through to Paraguay. Quoted by 
Aublet (1775) as being in French Guiana and by Schomburgk (1848) as in the Canuku 
Mountains. 

5. Selaginella pallescens (C. Presl) Spring in Martius, Fl. Bras. 1 (2) : 132 (1840); Alston in J. 

Bot., Lond. 72 : 226 (1934), Bull. Br. Mus. nat. Hist. (Bot.) 1 (8) : 231 (1955). Type from 

Mexico, Haenke s.n. (PR). 
Lycopodium pallescens C. Presl, Rel. Haenk. 1 : 79 (1825). Type as above. 
Lycopodium cuspidatum Link, Hort. Reg. Bot. Berol. 2:161 (1833). Type from Mexico. 
Selaginella cuspidata (Link) Link, Fil. Spec, in Hort. Reg. Bot. Berol. Cult. : 158 (1841). 

Spring in Mem. Acad. r. Sci. Lett. Belg. 24:66 (1850); Baker, Fern Allies 89 (1887); 

Hieron. in Hedwigia 58 : 286(1917). Type as above. 
Selaginella sulcangula Spring in Bull. Acad. r. Belg. 10 : 137 (1843), in Mem. Acad. r. Sci. 

Lett. Belg. 24 : 163 (1850), p.p. excl. planta jamaicensis. Type presumably from 

Venezuela. [Not collected by Plee in Colombia as stated by Spring.] 
Selaginella cordata Klotzsch in Linnaea 18 : 524 (1844), nom. superfl. (cites L. cordifolium 

Desv.) p.p. quoad spec. Moritz 67. 
? Selaginella cordifolia sensu Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 103 (1850), p.p. 

quoad Moritz 67. 
Selaginella emmeliana van Geert in Rev. Hort. Belg. Etrang. 10 : 220 (1884). Type from 

'Quinquina, South America'. 
Specimens seen: 

VENEZUELA. Without precise locality: Funck & Schlim 33 1 (BM). Aragua: below Guarnitas, Parque 
Nacional, 700 m, Alston 5859 (BM); Colonia Tovar, Moritz s.n. (NY); Fendler 321 (NY). Federal 
District: Venta Grande de Caracas, Moritz s.n. (NY); between Cotiza and los Venados, Allan 29 (NY, 
VEN); Between La Guayra and Caracas, Kuntze s.n. (NY); Silla de Caracas, Kuntze s.n. (NY); Cerro de 
El Avila, 1600 m, Pittier 29 (NY); Los Flores, Cerro de El Avila, 1600 m, Alston 5535 (BM); Galipan, 
Moritz 71 p.p. & 221 p.p. (BM); near Caracas, Ernst 212 (BM). Merida: Merida, Moritz 315 & 316 
(BM). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 253 

Miranda: Sebucan near Dos Caminos, Pittier 11918 (NY, VEN); Las Mostazas, between los Teques 
and Tejerias, 963 m, Allan 1 84 (VEN). 

Monagas: Cerro de la Cuevade Dona Anita, Steyermark 61881 (BM); Caripe, Moritz 68 (BM). 
COLOMBIA. Without precise locality: Moritz 69 (BM), 382 (BM). Magdalena: Santa Marta, Pur die 
s.n. (BM); near Santa Marta, Baker 6 (NY); Onaca Road, Santa Marta, 360 m, Smith 2243 (BM, NY, 
US); 7 ml E of Bonda, 600 m, Smith 2470 (NY). Valle del Cauca: Cartago, 1 950 m, Kuntze s.n. (NY). 

Geographical range: Mexico to Colombia. Alston (1936) records a specimen of this species 
from Brazil (Santa Catarina: Joinville, Schmalz 148 (NY)) which is now believed to be an 
escape from cultivation. 

Notes: It is not clear as to how Moritz collections became so mixed under especially nos 71 
and 22 1 (see page 272). 

6. Selaginella erythropus (Martius) Spring in Martius, Fl. Bras. 1 (2) : 125 (1840), in Bull 
Acad. r. Belg. 10 . 225 (1843), in Mem. Acad. r. Sci. Lett. Belg. 24 : 155 (1850), excl. var. 
major, Baker, Fern Allies : 103 (1887), excl. var. major, Knox in Trans, hot. Soc. Edinb. 
35 : 282 (1950). Type from Brazil : Piaui, Martius s.n. (BR, not seen). 

Lycopodium erythropus Martius, Icon. Select. PL Crypt. Bras. 39 (1834). Type as above. 
Selaginella setosa Linden, Cat. 23:9 (1869), Wochenschr. Vereines Beford. Gartenbaues 

Konigl. Preuss. Staaten 12 : 189 (1869). Type from Colombia. 
Selaginella erythropus var. minor Will. Watson in Gdnrs' Chron. Ill, 14 : 651 (1893) nom. 

nud.; Hieron. in Hedwigia 58 : 288 (1917). Type from 'Andes.' 
Selaginella pitcheriana Will. Watson in Gdnrs' Chron. Ill, 14: 651 (1893), nom. nud. in 

synonymy. 

Specimens seen: 

COLOMBIA. Caldas: Santa Cecilia, Pueblo Rico, 800 m, v. Sneidern 5053 (S). Cundinamarca: 

Tocarina, Lindig 1510 (BM). Tolima: Piedras, Andre s.n. (NY); R Saldana, 400 m, H aught 6379 (BM); 

Valle del Cauca: Chuchilla E. of Zarzal, 1 1-1 300 m, Pennell, Killip & Hazen 8504 (NY, US). 

ECUADOR. Guayas: Cerro de Soulana, Guayaquil, Jameson 585 (BM, US); Puna Is., Barclay 662 

(BM); Guayaquil, Eraser s.n. (BM). Manabi: Salanga, Barclay 652 (BM); Pinchincha: S Jose de Toachi, 

W of Quito, 915m, Bell 1 94 (BM). 

PERU. Ayacucho: Estrella, between Huanata and Rio Apurimac, 500 m, Killip & Smith 22664 (BM, 

NY). Junin: Colonia Perene, 600 m, Killip & Smith 24918 (BM, NY). Loreto: near Tarapato, Spruce 

3989 (BM, NY, US). San Martin: Juan Jui, Alto Rio Huallaga, 4-800 m, Klug 4267 (BM, COL); S bank 

of Perene River, within 1 5 m of confluence of Chanchamayo and Paucartambo, 670 m, Gascovne- Cecil 

6a (BM), 82a (BM); La Merced, Killip & Smith 23802 (BM, NY); Schunke Hacienda, above San Ramon, 

1 300-1 700 m, Schunke A 247 (BM); Chanchamayo Valley, Schunke 1 90 (BM). 

BOLIVIA. Beni: Secure, Guanay to Carahuarani, 548-914 m, Tate 1213 (US); Campolican, 

Rurrenabaque, Rusbv 764 (NY, US). La Plaz: San Buena Ventura, Williams 1399 (NY, US); Yungas, 

Rio Bopi Valley, Rushy 337 (NY, US). 

BRAZIL. Without exact locality: Glaziou 13364 (BM, C). Bahia: Fazend Barauna, 5 km S of Barreiras, 

Black 17877 (BM, IAN). Ceara: Bolland s.n. (K). Matto Grosso: Palmerras, Lindman A 2589 (BM). 

Minas Gerais: Near Diamantina, Moore 499 (BM); Macahuba, Silveira 394 (BM); between Sta Luzia 

do Rio das Velhas e Macaubas, Damazio 27384 (P); Lagoa Santa, Glaziou 22645 (G); Serra do Cipo, 

Damazio 1999 (BM). Piaui: without further locality, Gardner 2386 (M, P). Rio de Janeiro: St. Hilaire 

\62(P)\Hoehne92\ (BM). 

Geographical range: Colombia to Bolivia and east to the states of Piaui and Ceara in Brazil. 

7. Selaginella umbrosa Lemaire ex Hieron. in Engl. & Prantl, Nat. Pflanzenf. 1 (4) : 683 
(1901); in Hedwigia 58:287 (1917); Knox in Trans, bot. Soc. Edinb. 35:276 (1950); 
Alston in Bull. Br. Mus. nat. Hist. (Bot.) 1 : 30 (1952). Type from (?) Guatemala. 

Lycopodium umbrosum Lemaire ex Kuntze in Linnaea 23 : 292 (1850), nom. nud., in syn., 

non Bory ex Willd. ( 1 8 1 0). Type as above. 
Selaginella erythropus var. major Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 156 (1850); 

Baker, Fern Allies : 103 (1887). Type from Guatemala. 



254 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Selaginella lemairei Hieron. in Hedwigia 58 : 287 (1917), nom. provis. 

Specimens seen: 

COLOMBIA. Antioquia: Turbo, Schott 1 (NY); H aught 4800 (BM). 

Geographical range: Yucatan to Colombia; also in Tobago, where possibly introduced. 

8. Selanginella haematodes (Kunze) Spring in Martius, Fl. Brasil. 1 (2) : 126 (1840), in Mem. 

Acad. r. Sci. Lett. Belg. 24 : 156 (1850); Baker, Fern Allies : 103 (1887); Knox in Trans. 

hot. Soc. Edinb. 35 : 274 (1950); Alston in Bull. Br. Mus. nat. Hist. (Bot.) 1 : 234 (1955). 

Type from Peru, Loreto: Tocache Mission, Poeppig s.n. (K). 
Lycopodium haematodes Kunze in Linnaea 9 : 9 (1835), Farrnkr. 2:61 [excl. t.30 fig. C] 

( 1 843). Type as above. 
Selaginella filicina Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 189 (1850). Syntypes from 

Venezuela and Peru. 
Selaginella rosea Alston in J. Bot., Lond. 70 : 28 1 (1932). Type from Colombia. 

Specimens seen: 

VENEZUELA. Carabobo: San Esteban, near Puerto Cabello, 200 m, Alston 6109 (BM); Pittier 9234 
(NY, VEN). Federal District: Hacienda El Limon, valley of Puerto La Cruz, 500-1000 m, Pittier 9234 
(NY,VEN). 

COLOMBIA. Without precise locality, Triana s.n. (NY). Antioquia: Santa Elena, Archer 1 185 (BM, 
US); Ocateado, 2300-2600 m, Kalhreyer 1941 (K); El Penol, Daniel s.n. (BM); Las Isletas, Andre 434 
(NY); Villa Artega 3750-5000 m, Gutierrez & Barkley 17cl08 (BM, S). Cauca: Rio Huangubio, near 
Popayan, 1700-1800 m, Lehmann B T 875 (K, holotype of S. rosea Alston; NY, US). Choco: Truando 
Falls, Schott 7 (NY). Cundinamarca: near Bogota, Joseph A 465 (US); 1000 m, Triana s.n. (NY); 
Quebrada de Susumuco, Andre 963 (NY); Susumuco, 1000 m, Triana 696 (BM); La Palma, Triana s.n. 
(BM); Tequendama Falls, Joseph A 914 (BM); Paime, Joseph 1084 (BM). Meta: Villavicencio, Perez 
Arbelaez 1 (BM, US); 5000 m, Alston 7469 (BM); Rio Negro, S of Villavicencio, 457 m, Wright- 
Barker s.n. (BM); Plaza Bonita, Sierra Macarena, Philipson, Idrobo & Fernandez 1415 (BM); Sierra La 
Macarena, between Guapayita and 'Cano 20', Idrobo 4772 (BM). Narino: Putamayo upstream from 
Puerto Asis, Barclay 4754 (COL); Cordillera Oriental, Oreto, on Rio Oretopunqo, c. 450 m, Ewan 
16769 (BM). Santander: between Lebrija and San Vicente, 1400 m, Alston 7346 (BM). 
ECUADOR. Napo-Pastaza: Tena, Asplund 10188 (S); Mera, Asplund 18790 (S); between Puyo and 
Candos, 325-375 m, Mexia 6843 (BM); Tnte. H. Ortiz, 18 km from Puya on road to Tena, approx. 
1 km E of the road, 1 100 m, Ollgaard & Balsley 9232 (AAU). Santiago-Zamora: near Zamora, E of 
Loja, Camp E 50 (BM); between Rio Sabanilla and Canillones Tambo, 1370-2133 m, E of Loja, Camp 
E64(BM). 

PERU. Loreto: type as above; Tarapoto, Spruce 4036 (BM, NY); Puerto Arturo, 135 m, Killip & 
Smith 27747 (BM, NY); Lower R Huallaga, Williams 5359 (BM); Huanuco: near Riachuelo 
Chontalagua, Mexia 8302 (BM); Near Tingo Maria, Asplund 12072 (S), 12198 (S); 500-600 m, 
Ferreyra 1591 (BM), 2891 (BM), 2191 (BM); junction of Rio Monzon and Rio Huallaga, Asplund 
12693 (S). Junin: Rio Paucartambo Valley, near Perene Bridge, 700 m Killip & Smith 25292 (BM, 
NY); Cahupanas on Rio Pichis, 340 m, Killip & Smith 26776 (BM, NY); between Meriatiriani and 
Puero Yessup, Killip & Smith 26214 (BM, NY); Ucayali, Huber 1430 (BM); Perene River, within 
15 m of confluence ofChanchaymayo and Paucartambo Rivers, 1219 m, Gascoyne-Cecil 44 (BM). San 
Martin: Alto Rio Huallaga, Williams 6780 (BM). 

BOLIVIA. Without precise locality: Kelly s.n. (BM). Beni: Mapiri, 487 m, Williams 1397 (NY); 
1524 m, Rusby 450 (NY); Rio Chimate, 579 m, Tate 515 (BM); Charopampa near Mapiri, 570 m, 
Buchiien 623 (NY); San Antonio near Mapiri, Buchtien s.n. (Rosenst. exsicc. 94; BM). Chapare: Rio 
Juntas, Kuntze s.n. (NY). Cochamba: Espirito Santo, near Cochamba, Bang 1286 (BM, NY). La Paz: 
Polo-Polo near Coroico, prov. Yungas, 1100 m, Buchtien 624 (NY); Hacienda Sinaco, on way to 
Tipuani, 1400 m, Larecaja, Buchtien 5275 (BM); Bopi River Valley, Yungas mnts, 762 m, White 449 
(NY); 914m, Rusby 335 (NY), 377a (NY), 7 10 (NY); Tumapaca, Cardenas 1 974 (NY). 

Geographical range: Panama southwards to Bolivia. 

Notes: Alston (1932) described S. rosea from Colombia. We can see no difference between 
those specimens and pale forms of typical S. haematodes. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 255 

9. Selaginella coarctata Spring in Martius, Fl. Bras. 1 (2) : 126 (1840), in Mem. Acad. r. Sci. 

Lett. Belg. 24 : 164 (1850); Baker, Fern Allies: 101 (1887). Type from Brazil, Amazonas: 

Arara-Coara Mts, Rio Japura, Martius s.n. (M, fragment; BM). 
Selaginella puberula Klotzsch in Linnaea 18 : 522 (1844); Schomb., Reise. Brit. Guiana 3 : 

1042 (1848); Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 165 (1850); Baker, Fern Allies : 

101 (1887), p.p. excl. pi. peruviana; Knox in Trans, hot. Soc. Edinb. 35 : 254 (1950). Type 

from Brazil. 

Specimens seen: 

COLOMBIA. Vaupes: Raudal de Yurupari, Rio Vaupes, Schultes & Cabrera 19742 (US), 19748 (US); 

Cano del Caribe, Rio Guainia, 250-280 m, Schultes, Baker & Cabrera 1 8265 (US). 

BRAZIL. Amazonas: type as above; Rio Padauiri, Schomburgk 979 (BM, K, US); Campo on Rio 

Padauiri, Traill \4\7 (K). 

Geographical range: Eastern Colombia to north-west Brazil. Alston (1936) states 'British 
Guiana', assuming Schomburgk collected it on the nort-east side of Mt Roraima. In fact this 
reference to Roraima itself must be considered very doubtful, as the Schomburgk specimen 
(979) in K is labelled 4 Rio Padauiri' and his manuscript catalogue adds: 'grows in sandy 
savannahs'. It has never been recollected on Roraima; but Traill did get it again on the Rio 
Padauiri, and this is also the locality mentioned by J. Smith in /. Bot., Lond. 1 : 203 (1842). 

Notes: This species superficially resembles Selaginella asperula (No. 107), but the stems are 
pubescent, ex-articulate and terete (not glabrous, articulate and usually quadrangular), while 
the microspores have a plane (not convoluted) wing and larger warts. The pubescent 
branches separate S. coarctata from all other American species, except S. exaltata (No. 
104). S. puberula Klotzsch is based on a more luxuriant specimen, and is within the range of 
variation of S. coarctata. 

10. Selaginella longissima Baker in J. Bot, Lond. 19 : 208 (1881), 21 : 334 (1883), Fern 
Allies : 60 (1887); Hieron. in Bot. Jb. 34 : 580 (1905); Knox in Trans, bot. Soc. Edinb. 
35 : 289 (1950). Type from Colombia, Antioquia: Murri, 915m, Kalbreyer 1815 (K). 

Specimens seen: 

COLOMBIA. Antioquia: type as above. Cauca: Island of Gorgona, Cheesman 2a (BM). Choco: Alto de 

Buey, Nuqui, v. Sneidern A. 10 (S); Rio Nuqui, 200 m, Haught 5466 (BM, COL); between La Oveja 

and Quibdo, Archer 1743 (BM); La Concepcion, 15 km E of Quibdo, Archer 1964 (US); 1999 (US); 

Quibdo, Rio Atrato, 60 m, Archer 2025 (US). Narino: near Altoquer, 1000 m, Lehmann 85 (BM); 

Barbacoas, Alston 8520 (BM). Valle del Cauca: near Buenaventura, Alston 8663 (BM); Killip 11751 

(US), 11756 (US); Sabaletas, 25 m, Killip & Cuatrecasas 38786 (BM); near Cordova, Rio Dagua, 

80-100 m, Lehmann 1868 (BM); Tinier 525 (BM); Killip 5044 (NY, US), 5091 (BM, NY), 5253 (NY, 

US), 11781 (NY, US), 12154 (US). 

Geographical range: confined to Colombia and to lower elevations. 

Notes: In Killip 12 1 54 the costae of the leaves are thickened and blackened, perhaps because 
of a fungus disease. Similar states are known in other species, for example, S. novae- 
hollandiae (No. 45) and S. muscosa (No. 88). 

11. Selaginella mazaruniensis ['mazaruniense'] Jenman in Gdnrs' Chron. 22 : 2 10 (1897), 
Ferns Brit. W. Indies: 404 (1909). Type from Guyana, Essequibo: Mazaruni River, 
Jenman s.n. (NY, holotype; BM, isotype). 

Selaginella anceps sensu Wright in Trans. Linn. Soc. Lond. 6 : 88 (1901), non Lycopodium 

ancepsC. Presl(1825). 
Specimens seen: 

GUYANA. Essequibo: type as above; Potaro River, im Thurn 78 (K); Amatok, Potaro R, Jenman 
s.n. (BM, K, NY): Sandwith 1248 (BM, K): foot of the Kaieteur Falls, Potaro R, Jenman 1478 (K), s.n. 
(NY); Maguire & Fanshawe 23024 (BM), 23025 (BM): Macreba Falls, Kurufing R, Alison 342 (K): 
Mazaruni R, Jenman 7221 (K); Kaieteur Falls, Appun s.n. (BM): Tutin 518 (BM): Essequibo R, 
Jenman s.n. (BM): Roraima Range, 1066 m, McConnel & Quelch 706 (BM, K). 

Geographical range: Confined to the sandstone area of Guyana. 



256 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

12. Selaginella palmiformis Alston ex Crabbe & Jermy in Am. Fern. J. 63 : 141 (1973). 
Type from Venezuela, Amazonas: near Sake de Hua, in western foothills of Serra Imeri, 
800 m, Holt & Blake 490 (US, holotype; BM, isotype). 

Specimens seen: 

VENEZUELA. Amazonas: type as above; slopes of Mt Duida, 227 m, Tate 376 (NY). 

COLOMBIA. Amazonas: Rio Miritiparana 212 m, Schultes & Cabrera 16471 (BM, US). Vaupes: 

Cerro Isibukuri, Rio Kananari, 250-700 m, Schultes & Cabrera 14465 (US); Rio Guainia, Rio 

Naquieni, vicinity of Cerro Monachi, Schultes & Lopez 10104 (BM); Cano Teemeena, Rio Piriparana 

(tributary of R Apaporis), Schultes & Cabrera 17190 (COL, US), 17369 (COL, US), 17396 (US); Cano 

Oe-ge -dja, Jinogeje, Rio Apaporis, Schultes & Cabrera 1 7058 (BM, US). 

Geographical range: Confined to the Sierras of the Amazonian part of Venezuela and 
Colombia. 

13. Selaginella flabellata (L.) Spring in Flora, Jena 21 : 198 (1838), Bull. Acad. r. Belg. 10 : 
225 (1843); Baker, Fern Allies : 98 (1887); Alston in Bull. Br. Mus. nat. Hist. (Bot.) 1:31 
(1952). 

var. latifrons A. Braun in Annls Sci. nat. (Bot.) V, 3 : 278 (1865). Type from Venezuela, 

Merida: Colonia Tovar, Fendler 493 (BM). 
Selaginella mortoniana Alston MS, non Crabbe & Jermy. 

Specimens seen: 

VENEZUELA. Aragua: near El Portachuela, Parque Nacional, 1 100 m, Williams & Alston 172 (BM, 

US); Rancho Grande, Tschudi 38 (US). Merida: type, as above. 

Geographical range: Confined to the Cordiera Costanera of Venezuela. 

Notes: S. flabellata var. flabellata is confined to the Lesser Antilles. 

14. Selaginella mortoniana Crabbe & Jermy in Am. Fern J. 63 : 139 (1973). Type from 
Colombia, Tolima: near Mariquita, Linden 1004 (BM, holotype: BR, K, isotypes). 

Selaginella flabellata sensu Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 174 (1850), p.p. 

quoad pi. Linden 1004, non Lycop odium flabellatum L. (1753). 
Selaginella aciculata Alston MS. 
Selaginella hebetata Alston MS. 

Specimens seen: 

COLOMBIA. Without precise locality: Linden 1002 (BM); Funck & Schlim 57 (BM). Antioquia: 

Antioquia, Kalbreyer 2037 (BM). Boyaca: Emerald Mines, Muzo, 700 m, Lindig 1515 (BM). 

Cundinamarca: La Vega, 1200 m, Lindig s.n. (BM); La Palma, Lindig s.n. (BM); Cordillera de Bogota, 

Triana s.n. (BM). Tolima: type, as above. 

ECUADOR. Napo: at Rio Payamino, 60 km W of Coca, 350 m, Holm-Nielsen & Jeppeson 141 

(AAU); at Rio Suno 3 km W of Rio Napo, 400 m, Holm- Nielsen & Jeppeson 844 (AAU). 

Geographical range: Colombia and north-east Ecuador. 

15. Selaginella lechleri Hieron. in Engl. & Prantl, Nat. Pflanzenf. 1 (4) : 683 (1901), in 
Hedwigia 41 : 1 90 ( 1 902). Type from Peru, Puno: near San Gavan, Lechler 2 1 59 (BM). 

Specimens seen: 

COLOMBIA. Amazonas: Cachivera de Jirijirimo, Rio Apaporis, 250 m, Schultes & Cabrera 

12932 (COL, US), 14048 (COl, US); Soratama, Rio Apaporis, Schultes & Cabrera 16080 

(US); Cano Peritome, below Raudal Yayacopi, 227 m, Schultes & Cabrera 15526 (US). Vaupes: Cano 

Teemeena, Rio Piraparana, Schultes & Cabrera 17185 (COL, US); Alto Vaupes, between Plendaiva 

and Karuru, 400 m, Garcia- Bar riga 1 5009 (COL). 

PERU. Cuzco: between Mistiana and Keros, 700 m, Vargas 7372 (BM), 7393 (BM). Junin: 

Puerto Bermudez, 375 m, Killip & Smith 26467 (BM). Puno: type, as above. 

Geographical range: Colombia to south Peru. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 257 

16. Selaginella cheiromorpha Alston, sp. nov. (Figs 2 & 3) 

Species heterophylla ex affinitate S. quadrifariae sed foliis caulium minoribus et 

microsporis laevibus differt. 

Planta e basi repente erecta; caulibus glabris, usque ad 44 cm altis, basi 3 mm in diam.; 
parte inferiore simplici 4-23 cm longa, in siccitate straminea; rhizophoris basi restrictis, 
singulis vel binis; sobolis adeuntibus; parte frondosa ambitu late ovata, ramis usque ad 
2*5 cm inter se distantibus, bipinnatis ambitu deltoideis vel ellipticis, usque ad 1 5 cm longis 
et 12 cm latis. Folia caulium homomorpha distantiaque, oblongo-obovata serrulata, 
brevissime acuminata, 3 mm longa, 2*5 mm lata; folia lateralia horizontaliter patentia, 
plerumque leviter imbricata oblique ovato-oblonga, c. 6 mm longa, 3 mm lata, supra 
flavoviridia, subtus minus obscura; semifacie superiore semi-ovata, basi cordata, omnino 
serrata, serrulis partis basalis longioribus; semifacie inferiore semioblonga basin versus 
integra, apicem versus serrulata et cuneata; folia axillaria ovata, basi rotundato-cordata, 
4*5 mm longa, 2*5 mm lata, serrata; folia intermedia obovato-oblonga, 2/5 mm longa, 2 mm 
lata, imbricata, fere aequilateralia, sed obliqua, apice acuto, vel brevissime apiculato, 
ciliolata. Strobili in apicibus ramulorum singulatim dispositi, tetragoni, 1 cm longi, 
2*25 mm lati, nunquam ramosi; sporophylla ovato-deltoidea, carinata, serrata, apice 
brevissime apiculato; megasporae c. 360 jum crassae, subtiliter reticulatae; microsporae 
c. 24 jum crassae, pallide lutescentes, subtiliter granulatae. 

Typus: Colombia, Valle del Cauca : ad flumen Dagua, 950 m, Alston 7897 (BM, holotype). 

Other specimens seen: 

COLOMBIA. Valle del Cauca: valley of R. Dagua, 950 m Alston 780 IA (BM); 1050 m, Alston 7893 

(AAU, BM, COL, INPA, USM, US, VEN); La Cumbre, 1 800-2 1 00 m, Pennell 590 1 (NY, US). 

Geographical range: Confined to the western cordillera of Colombia. 

17. Selaginella terezoana Bautista in Bolm. Mus. Para. Emilio Goeldi (Bot.) II, 45 : 1 (1974). 
Type and sole specimen seen from Brazil, T. F. Roraima, '5°08'N— 60°40'W', Terezo 32 
(IAN). 

Geographical range: Only known from the type (above). The label cites coordinates which 
are in lowland Amazonas but it adds Tloresta Umbrofilia Sub-tropical,' suggesting montane 
forest. 

Note: The type specimen is sterile, but the very long cilia on the cauline and median leaves 
are most distinctive and like no other species. 

18. Selaginella anceps (C. Presl) C. Presl, in Abi Bohm. Ges. V, 3 : 581 (1844); Alston in J. 
Bot, Lond. 72:225 (1938); Knox in Trans, bot Soc. Edinb. 35:282 (1950). Type 
probably from Peru, Haenke s.n. (PR). 

Lycopodium anceps C. Presl, in Rel. Haenk. 1 : 80 (1825). Type as above. 

Lycopodium gracile Desv. ex Poiret, Encycl. Meth. Bot. Suppl. 3 : 551 (1814). Type from 

Peru. 
Selaginella gracilis (Desv.) Hieron. in Hedwigia 58 : 292 (1917), non Moore (1886). Type as 

above. 
Selaginella flab ellata sensu Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 174 (1850),p.p. pi. 

peruv. et var. expansa: Baker, Fern Allies : 98 (1887), p.p. quoad pi. andina, excl. syn. L. 
Selaginella casequiarensis Hieron. ex Bonap., Notes Pterid. 1 : 1 79 (19 1 5), nom. nud. 

Specimens seen: 

VENEZUELA. Bolivar: at the foot of Cerro Imei, Rio Pacimoni, Spruce 3380 (BM). 

COLOMBIA. Caqueta: Florentia, Quebrada de la Perdices, 400 mXuatrescasas 8851 (US). Valle del 

Cauca: Cordoba, 80-100 m, Killip 11811 (NY). 

ECUADOR. Napo-Pastaza: Tena, Asplund 8909 (S). 

PERU. Without precise locality or collector: Herb. Desvaux(P, type of L. gracile Desv.): type, as above. 

Ayacucho: Estrella between Huanta and Rio Apurimac, 500 m, Killip & Smith 2267 '1 (BM, NY). 



258 



A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 




IYPF SPECIMEN 

SEIAGINKI1A CHSIROilKfflA Alston 



Fig. 2 Selaginella cheiromorpha Alston: Type specimen, Alston 7897 (BM). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



259 




Fig. 3 Selaginella cheiromorpha Alston: A. Close-up of median leaves, x 16. B. Close-up 
showing lateral (/) and axillary (a) leaves, x 16. All from Alston 7897. 

Junin : Rio Pinedo, N of La Merced, 700-900 m, Killip & Smith 23605 (BM, NY); Rio Paucartambo 
Valley, near Perene Bridge, 700 m, Killip & Smith 25267 (BM, NY); along Rio Perene, near Colonia 
Perene, 600 m, Killip & Smith 25104 (BM, NY); on south bank of Perene River within 15 m from 
confluence of Rio Chanchosmayo and Rio Paucartambo, 670 m, Gascoyne- Cecil 6 (BM); Chanchos- 
mayo, Schunke s.n. (BM, P). Loreto: Tarapoto, Spruce 4053 (BM, NY); 750 m, Williams 6056 (BM); 
Creek Inche above Pongo de Manseriche, 215 m, Mexia 6371 (BM); Sierra del Pongo, 400 m, Mexia 
6185 (BM). San Martin: Juan Jui, Alto Rio Huallaga, 400 m, Klug 3831 (BM). Huanuco: Cochero, 
Dombeys.n. (BM); Tulmayo, near Tingo Maria, 6-700 m, Ferreyra 2161 (BM); near confluence of Rio 
Cayumba and Huallaga, 795-850 m, Mexia 8262 (BM); Pampayaco, Kanehira 1 18 (US). 
BOLIVIA. Without precise locality: Bridges s.n. (BM); Kelly s.n. (BM). La Paz: Isapuri, Laracaja, 
500 m, Williams 1398 (NY, US). 

Geographical range: Costa Rica southwards to Bolivia. 

Notes: The stem leaves of this species are almost uniform and directed upwards almost to the 
apex of the main stem, giving the plant a distinctive appearance. Presl's type in the 
herbarium of the National Museum at Praha is localized Luzon, but it was probably from 
Peru. S. casequiarensis Hieron. was so labelled on an herbarium sheet, but never validly 
published. Alston in MS was inclined to validate that taxon at specific rank, citing Spruce 
3380 (BM) as type. In our view the difference, i.e. the more pronounced cilia on the lateral 
leaves, is within the range of S. anceps. 

19. Selaginella hirsuta Alston ex Crabbe & Jermy in Am. Fern J. 63 : 138 (1973). Type from 
Venezuela, Bolivar: southeastern base of Carrao-tepui, 1460-1615 m, Steyermark 60844 
(US, holotype; BM, isotype) (the only specimen seen). 

Geographical range: Known only from south-east Venezuela. 

Note: Pubescence on the surface of the leaves is very rare in Selaginella. It is known in S. 
hispida (Willd.) Sw. from Jamaica, S. plagiochila Baker from Cuba, S. trachyphylla A. Br. 
from southern China, and S. tyleri (No. 52) from Venezuela. 



20. Selaginella bahiensis Spring in Martius, Fl. Bras. 
Bahia:£/artc/ze/2528(G, P). 



(2) : 124 (1840). Type from Brazil. 



260 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Two subspecies are recognised: 
Subsp. bahiensis 

Specimens seen: 

BRAZIL. Bahia: type as above; Blanchet s.n. (BM), 549 (G, NY); Soteropolis, Blanchet s.n. (G); 

Igregavelha, Blanchet s.n. (P); forests of Rio Grongoy basin, Curran 290 (B). 

Geographical range: Recorded only from eastern Brazil. 

Subsp. manausensis (Bautista) Jermy & Rankin, comb. nov. 

Basionym: Selaginella manausensis Bautista in Bolm Mus. Para. Emilio Goeldi (Bot.) II, 
45 : 2 (1974). Type from Brazil, Amazonas: along road from Manaus to Itacoatiara, at 
km 64, Rodrigues, Coelho & Monteiro 8588 (INPA, holotype; MG, isotype). 

Geographical range: Known only from the type. 

Notes: We could see only minor differences between Dr Bautista's plant and the type of S. 
bahiensis; the former has narrower, lanceolate-elliptic, denticulate, median leaves with an 
acute apex, compared with the obovate-elliptic, more coarsely toothed leaves, with an 
acuminate, even aristate apex, of subsp. bahiensis. We believe the two are closely related, but 
because of the geographical separation, accord them subspecific rank. 

21. Selaginella wolfii Sodiro, Crypt. Vase. Quito : 620 (1893). Type from Ecuador: woods of 
the western region, 1 800 m, Sodiro s.n. (P). 

Specimens seen: 

COLOMBIA. Choco: Bahia Solano, Haught 5525 (BM). Caldas: above Armenia, 1300— 1500 m, 

Pennell Killip & Hazen 8717 (BM, NY). Valle del Cauca: Queremal, Dagua, Perez 3207 (BM); 

1300 m, Alston 7907 (BM); Cordona, 80-100 m, Killip 5031 (NY, US), 5046 (NY, US); Cisneros, 

300-500 m, Killip 1 1489 (NY, US); valley of R Dagua, near sea level, Alston 8652 (BM); 900 m, Alston 

78 10 (BM); 950 m, Alston 7801 (BM); 1000 m, Alston 7896 (BM). Narino: Pambana, above Barbacoas, 

50m,£vva« 16842 (BM). 

ECUADOR. Without exact locality: type as above. Pastaza: southern border of Rio Napo, 2 km W of 

Yuralpa Holm-Nielsen & Jeppesen 995 (AAU). 

PERU. Cusco: In the region of Quincemil and Carratera from Cusco to Puerto Maldonado, Lobin s.n. 

(BM). 

Geographical range: Colombia and Ecuador, mainly on the western cordillera. The material 
recently collected in Peru is not typical, but we believe that it is within the range of the 
species. 

22. Selaginella hartii Hieron. in Urban, Symb. Antill. 3 : 525 (1903). Type from Trinidad: 
Tucuche, Crueger 181 (TRIN). 

Specimens seen: 

VENEZUELA. Sucre: peninsula of Paria, Cerro Patao, north of Puerto de Hierro, 850-860 m, 

Steyermark & Agostini 91202 (BM, US). 

Geographical range: Trinidad and Venezuela. 

23. Selaginella praestans Alston, nom. nov. 

Selaginella sprucei A. Braun in Annls Sci. Nat. (Bot.) V, 3 : 277 (1865), non Hook. (1861). 

Type from Peru, Loreto: woods of Rio Yacu-catina, Mt Guayrapurima, near Tarapoto, 

Spruce 4788 (G, holotype; BM, NY, isotypes). 
Selaginella anceps sensu Baker, Fern Allies : 104 (1887), p.p. non (C. Presl) C. Presl (1844). 

Specimens seen: 

COLOMBIA. Cauca: Chuare, Haught 5385 (BM, COL). 

ECUADOR. Napo-Pastaza: Mera, Asplund 18794 (S); banks of Rio Cotapino, c. 15 km W of Rac. 

Cotapino, 360 m, Whitmore 728 (BM). Pichincha: Rio Toachi, above confluence with Rio Pilaton, c. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 261 

915 m, Bell 248 (BM); Red-bark woods of Chimborazo, Spruce 5676 (BM); Archidona, Jameson 766 
p.p. (BM). 

PERU. Huanuco: Hacienda Mercedes, Cotirarda, 1500 m, Mexia 8196 (BM); Tingo Maria, Asplund 
12239 (S). Loreto: type, as above. 

Geographical range: Colombia to Peru. 

Notes: Braun cited the type of S. sprucei as Spruce 4780. This was a misprint, as has been 
ascertained from the type specimen at Geneve. 

24. Selaginella amazonica Spring in Martius, Fl. Bras. 1 (2) : 124 (1840), in Mem. Acad. r. 
Sci. Lett. Belg. 24 : 176 (1850); Baker, Fern Allies : 103 (1887). Type from Brazil; without 
precise locality, Martius s.n. (M). 

Selaginella rionegrensis Baker in J. Bot., Lond. 23 : 117 (1885), Fern Allies : 101 (1887). 
Type from Brazil. 

Specimens seen: 

VENEZUELA. Amazonas: Cerro de la Neblina, Rio Yatua, between camps 2 and 3, 200-600 m, 

Maguire, Wurdack & Bunting 37393 (BM, NY). 

COLOMBIA. Amazonas: Raudal de Jirijirimo, R Apoporis, Schultes 12083 (BM); Garcia 13726 (BM). 

Vaupes: near Piedra de Cocui, Rio Negro, Schultes & Lopez 9523 (BM); Raudal de Jirijirimo, Rio 

Apoporis, 250 m, Schultes & Cabrera 12458 (COL, US); Cano Paca, Raudal Yo-ree-hwe-ro, Rio 

Piriparana, Schultes & Cabrera 17562 (US), 17570 (US); Cachivera Miriti, 250 m, Schultes & Cabrera 

14410 (COL, US), 14410c (US); Rio Vaupes, Javarete, Schultes & Cabrera 19422 (UC). 

BRAZIL. Without precise locality: type as above. Amazonas: S of Serra Cuputy, Rio Uyapura, Herb. 

Capanema 32764 (BM); near Panure, Rio Uaupes, Spruce 2501 (BM, type of S. rionegrensis Baker); 

base of Serra Tukano, above Trovao, Rio Uaupes, Schultes & Pires 9005 (BM); Varadouro, Uaupes, 

Luetzelburg 23646 (M); San Gabriel, Cuabrugas, Spruce 2358 (K); San Gabriel, Spruce 2053 (K); 

Estrada Campos Salles, Manaos, Huebner 67 (B). 

Geographical range: Venezuela, Amazonian Brazil and Colombia. 

25. Selaginella oaxacana Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 177 (1850). Type from 
Mexico, Oaxacana: Chinanbla, Galeottii 6808 bis (BM, P). 

Selaginella anceps sensu Baker, Fern Allies : 104 (1887); sensu Christ in Pittier, Primitiae 
Florae Costaricensis 3 (1) : 61 (1901); non Lycopodium anceps C. Presl. 

Specimens seen: 

COLOMBIA. Choco: La Concepcion, 15 km E of Quibdo, 75 m, Archer 1974 (BM). Cundinamarca: 

Andes of Bogota, Triana s.n. (P). Narino: Tuquerres road, Barbacoas, 900 m, Triana 696 (BM, COL). 

Valle del Cauca: Cordoba, Killip 5092 (US); v. Sneidern 4467 (S); Costa del Pacifico, Rio Raposo, 

20-50 mjdrobo 5255 (BM). 

ECUADOR. Napo-Pastaza: Mera, to approx. 4 km N of the village (along Rio San Jorge and Rio Tigre) 

1200 m. Ollgaard & Balslev 9074 (AAU, BM). Santiago-Zamora: Cordillera Cutucu, Camp E 1 194 

(BM). 

Geographical range: Mexico to Ecuador. 

Notes: Alston placed South American material in this species with some reservation. We 
feel, with the material available, there is no good reason for distinguishing it as a separate 
taxon. 

26. Selaginella quadrifaria Alston, sp. nov. (Figs 4 & 5) 

Species heterophylla ex affinitate S. praestantis, sed foliis intermediis minus inaequilateral- 
ibus, foliisque caulium apicem versus magnis, quadrifariis, et magnitudine fere aequalibus, 
differt. 

Planta e basi breviter repente erecta; caulibus glabris usque ad 55 cm altis, basi 3 mm in 
diam; parte inferiore simplici c. 32 cm longa, in siccitate pallide straminea, subdistantibus 
tecta superiorem partem versus foliis majoribus minus aequalibus; rhizophoris basi restrictis; 
parte frondosa ambitu late ovata, ramis usque ad 4 cm inter se distantibus, bipinnatis, 



262 



A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 




PLANTS OF PERU 1 2 OlC 53 

re to 



VwMm^ G2S2 



TYPE SPECIMEN 

3EIACIKEUA QUMEIPiHIA Alaton 



Fig. 4 Selaginella quadrifaria Alston: Type specimen, Mexia 6282 (BM). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



263 




Fig. 5 Selaginella quadrifaria Alston: A. Close-up of lateral (/) and axillary (a) leaves, x8. B. 
Close-up of median leaves, x 8. C. Lateral leaf showing acroscopic auricle, x 32. D. Apex of 
lateral leaf, x224. All from Mexia 6282. 

ambitu ovatis vel deltoideis, usque ad 20 cm longis, 12 cm latis. Folia caulium in parte 
infima deltoidea, integra, subacuta, 2' 5 mm longa, in parte superiore ovata, usque ad 6 mm, 
serrulata, basi nonnunquam ciliata, apice leviter acuminata; folia lateralia partis frondosae 
fere horizontaliter patentia, leviter imbricata, oblonga, c.8 mm longa, 3 mm lata, fere 
concoloria; semifacie superiore semi-oblonga, basi leviter cordata, et ciliata, alibi serrulata; 
semifacie inferiore semi-oblonga, basi retracta, attenuata et excavata, integra; folia axillaria 
oblongo-deltoidea, basi utrinque cordata et ciliata; folia intermedia oblongo-ovata, 3*5 mm 
longa, 3 mm lata, divergentia, fere aequilateralia, vix acuminata. Strobili in apicibus 
ramulorum singuli vel bini dispositi, tetragoni, 1*5 cm longi, 2 mm lati, nonnunquam 
ramosi; sporophylla ovato-deltoidea, carinata, serrata, sensim acuminata; megasporae c. 
400 /um albidae, subtiliter reticulatae; microsporae c. 27 jum crassae, pallide lutescentes, 
papillis elongatis crebre obsitae. 

Typus: Peru, Loreto: Sierra de Pongo, 500 m, legit Mexia 6282 (BM, holotype). 

Other specimens seen: 

COLOMBIA. Without exact locality: 'Southern', Hopp s.n. (B, fragm. BM). 

PERU. Junin: Puerto Yessup, 400 m, Killip & Smith 26372 (BM, NY). 

Geographical range: Southern Colombia to northern Peru. 



27. Selaginella tanyclada Alston ex Crabbe & Jermy in Am. Fern J. 63 : 143 (1973). Type 
from Colombia, Choco: La Concepcion, 15 km E of Quibdo, 75 m, Archer 1966 (US, 
holotype; BM, isotype). 



264 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Specimens seen: 

COLOMBIA. Choco: type, as above; Alto de Buey, Nuqui, 1000 m, Sneidern s.n. (S). 

Geographical range: Confined to Colombia. 

28. Selaginella bombycina Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 191 (1850); Baker, 
Fern Allies : 80 (1887); Knox in Trans, bot. Soc. Edinb. 35 : 284 (1950); Alston in Bull. Br. 
Mus. nat. Hist. (Bot.) 1:236 (1955). Type from Peru, San Martin: Juana del Rio, 
Matthews \1S\ (K). 

Selaginella wallisii Regel ex Salom., Nomencl. : 358 (1883), nom. nud. 
Selaginella anceps sensu Sodiro, Crypt. Vase. Quit. : 621 (1893), non C. Presl. 

Specimens seen: 

COLOMBIA. Antioquia: Sonson, Rioverde, Rio Verde de los Montes, Gutierrez 35629 (UC). 

Narino: Costa del Pacifico, Rio Telembi, Barbacoas, 30 m, Idrobo & Weber 1459 (BM). Tolima: k La 

Trinidad\ Libano, 1 100-1300 m, Pennell 3314 (NY, US). Valle del Cauca: bank of R Engano, Dagua 

Valley, 700 m, Alston 7859 (BM); Cisneros, 300-500 m, Killip 35543 (COL). 

ECUADOR. Esmeraldas: Playa Rica, Parroquia de Concepcion, 105 m, Mexia 8434a (BM), 8475 

(BM); Angamarca, Leon, Sodiro s.n. (NY). Guyas: near Quillallpa, 150 m, Fagerlind & Wiborn 679 

(S); Teresita, 3 km W of Bucay, 270 m, Hitchcock 20484 (BN, NY, US); Tenguel Estate, near 

Guayaquil, Haught 2876 (BM); near Bucay, 300-375 m, Camp E3847 (BM). La Paz: Polo-Polo near 

Coroico, Nordyungas, 1 100 m, Buchtien 3553 (US). Los Rios: Hacienda Clementina, Cerro Mombe, 

600 m, Asplund 5551 (S) Pastaza: road Puyo-Macas, 31 km from Puyo, 1 100 m, Ollgaard & Balslev 

9026 (AAU). Santiago-Zamora: near Chupiantza, lower plain of R Upano, 500-700 m, Camp El 022 

(BM). 

PERU. San Martin: type, as above; near Tingo Maria, Allard 20416 (BM), 20442 (BM), 20834 (BM), 

22194 (BM). 

Geographical range: Costa Rica to Peru. 

Notes: Ollgaard & Balslev 9026 is a form with most median leaves having a short apical 
arista; those on the main stem, however, show the more typical aristate leaves. Specimens 
from Costa Rica (Standley 37269) have been identified as S. wallisii Regel by O. C. Schmidt; 
it seems probable, therefore, that this name belongs here. 

29. Selaginella radiata (Aublet) Spring in Bull. Acad. r. Belg. 10 : 143 (1843), in Mem. Acad. 
r. Sci. Lett. Belg. 24 : 120 (1850), p.p. excl. pi. orbigniana; Alston in Reprium Spec. nov. 
Regni veg. 40 : 308 (1936), in Pulle, Fl. Surin. 1 : 165 (1938). Type from French Guiana, 
without precise locality, Aublet s.n. (BM). 

Lycopodium radiatum Aublet, Hist. Pi Guian. 2 : 967 (1775), p.p. excl. syn. Dillen. Type as 

above. 
Lycopodium penniforme Lam., Encycl. 3 : 20 ( 1 79 1 ), excl. var. /?. Type from French Guiana. 
Stachygynandrum penniforme (Lam.) Beauv., Prodr. Aethog. : 1 10 (1805). Type as above. 
? Selaginella descrescens Spring in Van Heurck, PI. Nov. : 28 (1 870). Type from Brazil. 
Selaginella penniformis (Lam.) Hieron. in Hedwigia 58 : 295 (1917). Type as above. 
Selaginella haenkeana sensu Spring Mem. Acad. r. Sci. Lett. Belg. 24: 187 (1850), p.p. 

quoad pi. guian.; Baker, Fern Allies : 102 (1887), p.p. excl. syn. S. dimorpha Klotzsch. 

Specimens seen: 

FRENCH GUIANA. Without precise locality: type as above; Herb. Lamarck s.n. (P, type of L. 

penniforme): Karouany, Sagot 838 (BM, NY). 

SURINAM. Without precise locality: Focke 1254 (U). Nickerie: Upper Nickerie R, Tulleken 333 (L), 

5 10 (L). Suriname: Surinam R, Tresling24\ (U). 

GUYANA. Essequibo: near Chodikar landing, Acarai Region, Guppy 457 (BM); Manicole bog, near 

Onoro River, Guppy 336 (BM). 

COLOMBIA. Cundinamarca: Cordillera Oriental, between Sebastopol and Analaima, 2285 m, Little 

8580 (COL); Cordillera Oriental, at railroad station Tablanca, 40 km W, NW of Bogota, c. 2200 m, 

Little 9\55 (COL). 

BRAZIL. Without exactly locality: Glaziou 10213 (C). Matto Grosso: St. Manoel, Hoehne 5275 (BM). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 265 

Para: Belterra, Black 41-916 (BM); Fabrica mata de Najatenua, Rio Moju, Black 16330 (BM, IAN, U); 
Rio Cumina, Sampaio 5039 (BM); Barra do Rio Jurnena, Pires 3679 (BM); St Antonio de Prata, Huber 
7044 (BM); Rio Trombetas, near Lake Orapecu, Traill 1418 (K); Rio Vermelha, Grute Verde, 
Tocantima; Froes 2701 1 (BM, IAN); Igarpe Pontanarri, Rio Oiapoque, Froes 26007, (BM, IAN). 

Geographical range: The Guianas, Colombia and N Brazil. 

Notes: A very small but distinct plant from Brazil, Santarem, Spruce. 682 and 943 (BM, 
CGE, GH: syntypes of S. decrescens Spring) was considered by Alston (in sched.) to be a 
depauperate form of this species. On further investigation it may turn out to be a distinct 
species. Aublet quotes 'Lycopodioides radiatum, fdicinum, minus et angustius Dillen., 
Muse. p. 470, t. 67, no. 7', by which he probably intended 'Lycopodioides denticulatum 
erectum filicinum, minus et argutius Dillen., Muse. p. 470, t. 65, no. 7\ which was from 
China and represents S. moellendorffii Hieron. (specimen in Oxford (OXF) has been deter- 
mined by Alston). The name appears to have been confused with Lycopodioides radiatum 
dichotomum Dillen., Muse. p. 474, t. 67, a true Lycopodium. It seems best therefore to follow 
Baker and accept Aublet's specimen, which Baker unfortunately misidentified, as the type. 

30. Selaginella speciosa A. Braun in Annls Sci. nat. (Bot.) V, 3 : 273 (1865); Knox in Trans, 
bot. Soc. Edinb. 35 : 289 (1950). Type from Colombia, Cundinamarca: Bogota, Cuervo 
[herb. Triana] s.n. (BM). 

Selaginella huberi Christ in Bull. Herb. Boissier II, 1 : 73 (1901); Hieron. in Hedwigia 43 : 6 
(1904). Type from Peru. 

Specimens seen: 

COLOMBIA. Without precise locality: Wallis s.n. (US). Amazonas: Arriba de la desembocadura del 

rio Loretoyacu, Duque-Jaramillo 2333 (COL). Antioquia: without further locality, Kalbreyer s.n. 

(BM); Quebrada San Julian, between Argelia and El Tigre, 2760-2900 m, Ewan 15778 (BM). Caldas: 

Quebrada Pontona, 1 8 km W of La Dorada, 400 m, Haught 2 1 32 (BM). Cundinamarca: type as above. 

ECUADOR. Napo-Pastaza: near Canelos, 3-400 m, Mexia 6869 (BM); in sylvis Archedoniae, 

Jameson 766 p.p. (BM). Santiago-Zamora: near Mendez, 425-750 m, Camp E892 (BM). 

PERU. Loreto: near Tarapoto, Spruce 4628 (BM, CGE, NY); between Yurimaguas and Balsapuerto, 

135-150 m, Killip & Smith 28340 (BM, NY); Balsapuerto, 220 m, Klug 2921 (BM); Balsapuerto, 

150-350 m, Killip & Smith 28403 (BM, NY); Sierra del Pongo, 600 m, Mexia 6283 (BM); River 

Huallaga, Huber 1 547 (P, type of S. huberi Christ). 

Geographical range: From Colombia to N Peru. 

31. Selaginella haenkeana Spring in Bull. Acad. r. Belg. 10 : 225 (1843); in Mem. Acad. r. 
Sci. Lett. Belg. 24 : 187 (1850), p.p.; Hieron. in Hedwigia 58 : 295 (1917); Alston in J. Bot., 
Lond. 72:226 (1934). Type from Peru, k Cordilleris Chilensibus', Haenke s.n. (PR, 
holotype; B, US, isotypes). 

Lycopodium plumosum sensu C. Presl, Rel. Haenk. : 79 (1825), non L. (1753). 

Selaginella dimorpha Klotzsch in Linnaea 18 : 523 (1844). Type as above. 

Selaginella leptoblepharis A. Braun in Annls Sci. nat (Bot.) V, 3 : 279 (1865). Syntypes from 

Colombia. 
Selaginella hartwegiana var. leptoblepharis (A. Braun) Baker in J. Bot., Lond. 23: 118 

(1885), Fern Allies: 102(1887). 

Specimens seen: 

COLOMBIA. Caldas: Rio Santa Rita, Salento, 16-1800 m, Killip & Hazen 8958 (NY). Cundinamarca: 
Salto de Tequendama, 2100 m, Triana s.n. (BM, syntype collection of S. leptoblepharis A. Braun); 
Lindig 1506 (BM); Haught 6404 (BM); Holton 83 (NY); Alston 7416 (BM), 7423 (BM); Joseph s.n. 
(US); La Vega, 2400 m, Lindig s.n. (BM, syntype collection of S. leptoblepharis A. Braun); between La 
Vega and Facatatioa, 2200 m, Haught 6136 (BM); Alban, 1600 m, Guevara 305 (US); Cordillera 
Oriental, falls of Rio Bogota, vicinity of El Salto, c. 2500 m, Little 7879 (COL); Villete, 800 m Idrobo & 
Dumont 4549 (BM); Salto de Tequendama, 2500 m, Cuatrecasas 78 (COL). Santander: Corcoba, near 
Bucaramanga, 2200 m, Alston 7319 (BM). Tolima: k La Trinidad', Libano 1 100-1300 m, Pennell 



266 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

3314A (NY); Ibague; corregimiento Juntas, finca El Silencio, 2800 m, Echeverri 869 (COL). Valle del 

Cauca: valley of the R Digua, 950 m, Alston 7834 (BM); 1050 m, Alston 7894 (BM); near Queremal, 

1300 m, Alston 7905 (BM); banks of Engano, Digua Valley, 700 m, Alston 7862 (BM); 'La Gallera', 

Micay valley, 1 400-1 500 m, Killip 7712 (NY, US), 7765 (NY, US). 

ECUADOR. Azuay: Chacanceo, Loma de la Plata, 1430-1465 m, Steyermark 52683 (BM); near 

Portovelo, Rose 23450 (NY). Napo: Baeza, c 1 km SW of the village 2000 m, Ollgaard & Balslev 

10215 (AAU). 

PERU. Without exact location: type as above; Matthews 18 (BM). Junin: Pichis Trail, Dos de Mayo, 

1700-1900 m, Killip & Smith 25843 (BM, NY); Schunke Hacienda, above San Ramon, 1300-1700 m, 

SehunkeA24% (BM,US). 

BOLIVIA. Laguna: In antris humidis montium Nuevo Mundo, Orbigny s.n. (P). 

Geographical range: Colombia, Ecuador, Peru and Bolivia. 

Notes: PresFs type in the National Museum at Praha was labelled 'Cordilleras de Chili', but 
no doubt Haenke collected it in Peru. Selaginella dimorpha Klotzsch is an exact synonym of 
S. haenkeana as both were based on Lycopodium plumosum sensu Presl. 

32. Selaginella hartwegiana Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 188 (1850); Baker, 
Fern Allies : 102 (1887); Knox in Trans, hot. Soc. Edinb. 35 : 256 (1950). Syntypes from 
Ecuador, Pinchincha: in declivitate Andium prope Nanegal, Hartweg 1477 (BM, LG); 
Tungurahua: in sylvis prope Banos, Jameson s.n. (K); ad Pillzhum, Jameson s.n. (K). 

Selaginella hartwegi Spring ex Benth., PI. Hartweg. : 359 (1857), nom. nud. based on 

Hartweg 1477. 
Selaginella lizarzaburui Sodiro, Crypt. Vase. Qui. : 615 (1893). Type from Ecuador. 

Specimens seen: 

COLOMBIA. Cundinamarca: Villete, 800 m, Idrobo & Dumont 4549 (BM). Narino: near Ricuarte, 
1 500 m, Alston 8434 (BM). Valle del Cauca: near Queremal, 1400 m, Alston 7923 (BM). 
ECUADOR. Without precise locality: Jameson 27 (NY), 106 (BM). Chimborazo: in red-bark woods, 
Spruce 5674 (BM, CGE); near Huigra, Rose 23962 (NY); near Las Juntas, Rose 23177 (US). Napo- 
Pastaza: Mera, Asplund 19677 (S); Mera, road and mule tracks to c. 4 km N of the village, 1200 m, 
Ollgaard & Balslev 9118 (AAU, BM); Baeza-Tena 8 km from Baeza, 1 900—2000 m, Balslev & Madsen 
10368 (AAU); road Baeza-Lago Agrio, Rio Salado, 49 km from Baeza, 1400 m, Balslev & Madsen 
10280 (AAU, BM). Pichincha: type as above; Saloya, 1800 m, Acosta Solis 5626 (BM); El Volante, 
between Chiriboga and Santo Domingo de los Colorados, Asplund 1 7386 (S); between Quito and Santo 
Domingo de los Colorados, 1450 m, Holdridge 1588 (BM); near Rio Negro, Pastaza Valley, 1250 m, 
Diels 914 (B); Mt Atacazo, near St Florencio, 1600 m, Sodiro s.n. (P, type of S. lizarzaburui Sodiro); St 
Florencio, 1600 m Andre 3632 (NY); Guyas, Hacienda Solento, near Santa Rosa, Canton Pajili, Leon, 
1000 m, Mexia 6723 (BM); Tungurahua: syntypes as above; between Machai and La Victoria, Rio 
Pastaza, Asplund 8508 (S); Banos, 2000-2400 m, Lehmann 361a (BM); between Banos and Cashurco, 
1 300-1 800 m, Hitchcock 2 1 869 (NY, US); 2500 m, Lehmann 383 (BM). 

Geographical range: Colombia and Ecuador. 

Notes: Selaginella haenkeana (No. 31) is the closest ally, but has more obtuse and more 
oblong lateral leaves. Selaginella hartwegiana superficially resembles S. pearcei (No. 34), 
which has denticulate (not ciliate) lateral leaves and usually acuminate (not aristate) median 
leaves. Selaginella novae- hollandiae (No. 45) is a smaller species, branched from near the 
base, with the general outline oblong rather than ovate. 

33. Selaginella viticulosa Klotzsch in Linnaea 18 : 524 (1844); Spring in Mem. Acad. r. Sci. 
Lett. Belg. 24 : 186 (1850); Baker, Fern Allies : 102 (1887); Knox in Trans, bot. Soc. Edinb. 
35 : 276 ( 1 950). Type from Venezuela, Federal District: near La Guayra, Moritz 70 (BM). 

Specimens seen: 

VENEZUELA. Without precise locality: Karsten s.n. (BM); Funck 344 (BM); Fendler 258 (NY). 
Carabobo: Lomas de Turismo, 100-200 m, Pittier 13866 (B); San Esteban, near Puerto Cabello, Alston 
6114 (BM), 6130 (BM); Hacienda Marture, between La Entrada and Las Trincheras, 500 m. Williams 



SELAGINELLA IN TROPICAL SOUTH AMERICA 267 

& Alston 352 (BM). Federal District: type as above; between Caracas and La Guayra, Rose 2 1 725 (NY); 

Galipan, Funck 274 (P). 

COLOMBIA. Without exact locality: herb. A. Braun s.n. (BM). 

Geographical range: Costa Rica and Panama to Colombia and Venezuela. 

Notes: Alston collected this species on a wall at Tijuca near Rio de Janeiro in 1938 and 
believed it to be an escape from cultivation. It is also said to be introduced into Trinidad and 
Bermuda. The specimen from Colombia without locality (ex hort. Berol. et hb. A. Braun) is a 
young specimen which lacks a fully developed frond branch system. 

34. Selaginella pearcei Baker in /. BoL, Lond. 22 : 246 (1884), Fern Allies : 75 (1887). Type 
from Peru, Huanuco: Cordilleras of Pozuzo, Pearce 249 (K). 

Selaginella oligoclada Baker in /. Bot., Lond. 22 : 277 (1884). Type from Colombia. 

Specimens seen: 

VENEZUELA. Merida: La Mucuy, Vareschi & Pannier 1373 (BM); Sierra Nevada, above Merida, 

2800 m, Alston 6779 (BM). Trujillo : above San Rafael de Bocono, 2500 m, Box 3830 (BM). 

COLOMBIA. Boyaca: around Bocota, c. 2200 m, Grubb & Guvmer P83 (COL). Cauca: above 

Carpinteria 2500 m, Alston 8217 (BM); W of El Tambo, 2500 m, Haught 5209 (BM, COL); 

Magdalena: near Playoncito, Santa Marta, Schultze 1518 (B, BM). Norte de Santander: near Ocana, 

24-2700 m, Kalbrever 1080 (K, type of S. oligoclada Baker). Santander: near La Baja, 22-2300 m, 

Killip & Smith 1 8280 (NY); 3000 m, Killip & Smith 1 8348 (NY). 

PERU. Without precise locality: Matthews s.n. (BM). Huanuco: type as above. Junin: between S Jose 

and La Achira, Montanas de Pangoa, Raimondi 1 563 (B, BM). 

Geographical range: Venezuela to Peru. 

Notes: Baker (1884) described a specimen from the Andes of Ocana, Colombia (Kalbreyer 
1080; K) as S. oligoclada. This is a more robust plant with less pinnate branching and with 
median leaves slightly broader with a distinct cusp. Intermediate forms with S. pearcei are 
found, and it is doubtful if the former species can be maintained. S. pearcei may be confused 
with depauperate non-flagellate specimens of S. flagellata (No. 41), but the latter has more 
attenuate, almost aristate, median leaves. 

35. Selaginella meridensis Alston, sp. nov. (Fig. 6). 

Species heterophylla ex affinitate S. viticulosae sed foliis intermediis nee ciliatis nee 
auriculatis differt. 

Planta e basi breviter repente erecta; caulibus glabris, circa 23 cm altis, basi 1*5 mm in 
diam.; parte inferiore simplici c. 9 cm longa, in siccitate pallide virido-straminea, foliis fere 
ad basin heteromorphis; rhizophoris ad partem basin versus restrictis; parte frondosa ambitu 
oblonga, ramis usque ad 2*5 cm inter se distantibus, bipinnatis, ambitu irregularibus, usque 
ad 7 cm longis, 3 cm latis. Folia lateralia patentia, fere contigua, oblonga, 5 mm longa, 2 mm 
lata, subacuta; semifacie superiore semi-oblongo-lanceolata, basi gradatim rotundata, crebre 
serrata vel plusminusve ciliolata, apicem versus distanter serrata; semifacie inferiore semi- 
oblonga, basi leviter rotundo-cuneata et ciliolis nonnullis instructa, alibi integra; folia 
axillaria oblongo-lanceolata, serrata, 3'5 mm longa, basi utrinque gradatis rotundata; folia 
intermedia ovato-oblonga, 1*5 mm longa, 1mm lata imbricata, longe aristata. Strobili 
plerumque in apicibus ramulorum brevium lateralium singuli vel bini dispositi, tetragoni, 
3 mm longi, L75mm lati, non ramosi; sporophylla deltoidea, carinata, serrulata, vix 
acuminata; megasporae c. 380, pallide flavido-brunneae (humectae), subtiliter reticulatae; 
microsporae papillosae. 

Typus: Venezuela, Merida: supra Las Cadras, ad septentriones Torondov, 1820-2255 m, 27 
Mar. 1944, Steyermark 55791 (BM, holotype; US, isotype). 

Other specimens seen: 

ECUADOR. Azuay: between Cruz Pamba and Loma de Canela, 2315-2500 m, Steyermark 52967 

(BM, US). Median, lateral and axillary leaves all lack cilia but in other characters resemble the type. 



268 



A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 




Fnm THE ONITSO STATES NATIONAL HERlAStU 

SaLaginella 



Venezuela: Above hut C adras, north of 

Torondor, Uerlda, 1**0-2255 ffi. 
J. A. StayenaarV 55791 Wr. 27, 1944 



TYPE SPECIMEN 

SEIAGIKEUA JrERIDENSIS Alston 



Fig. 6 Selaginella meridensis Alston: A. Type specimen, Steyermark 5579 1 (BM). B. Close-up of 
lateral (/) and axillary (a) leaves, x 16. C. Close-up of median leaves, x 33. All from Steyermark 



^7Q1 



SELAGINELLA IN TROPICAL SOUTH AMERICA 269 

Geographical range: Venezuela to Ecuador. 

36. Selaginella popayanensis Hieron. in Hedwigia 43:9 (1904). Type from Colombia, 
Cauca: near Popayan, Lehmann 6968 (US, holotype; K, isotype). 

Selaginella moritziana var. pseudapoda A. Braun in Annls Sci. nat. (Bot.) V, 3 : 285 (1865). 
Type from Colombia. 

Specimens seen: 

COLOMBIA. Cauca: type as above; Macizo Colombiano, Las Papas paramo, between El 

Boqueron and La Hoyola, 3200-3510 m, Idrobo, Pinto & Bischler 3516 (BM); Purace, km 6, 2750 m, 

Alston 8063 (BM). Cundinamarca: Manzanos, 1700 m, Lindig 1518 (BM). Santander: Pamplona, 

Joseph s.n. (BM, US). Valle del Cauca: near Queremal, 1 300 m, Alston 7906 (BM). 

ECUADOR. Carchi: Volcan de Chile, near Tufino, 3630 m, Wiggins 10637 (BM). Pichincha: Valley 

ofLloa, Benoist 2700 (P). 

BOLIVIA. Without precise locality: Bang 80 (BM), 1 1 1 (BM), 1 29 (BM). La Paz: Yungas. Jay c (NY), e 

(NY); 1 830 m, Rusby 462 A (NY). 

Geographical range: Colombia to Bolivia. 

Notes: As delimited above this species is very variable, and may need further subdivision. S. 
cavifolia (No. 37) differs by having stems which bear rhizophores throughout and relatively 
broader lateral leaves; S. jlacca (No. 38) also bears rhizophores throughout. S. moritziana 
(No. 42) bears rhizophores only at the base, but is branched there and is more closely leafy. S. 
cruegeri Jenman from Trinidad is a little known species of similar habit, but with the lateral 
leaves distinctly ciliolate at the base. 

37. Selaginella cavifolia A. Braun in Annls Sci. nat. (Bot.) V, 3 : 272 (1865); Baker, Fern 
Allies: 43 (1887). Syntypes from Colombia, Cundinamarca: Salto de Tequendanna, 
Triana s.n. (BM); Boqueron, Bogota, 2700 m, Lindig 1511 (BM). 

Selaginella moritziana var. elongata A. Braun in Annls Sci. nat. (Bot.) V, 3 : 285 (1865). 
Syntypes from Venezuela and Colombia. 

Specimens seen: 

VENEZUELA. Merida: Tovar, Fendler 322, 323 (BM; syntypes of S. moritziana var. elongata A. 

Braun). 

COLOMBIA. Cundinamarca: syntypes as above; Salto de Tequendama, 2100 m, Alston 7413 (BM); 

Bogota, Rio Arz, 2900 m, Lindig 1505 (BM; syntype of S. moritziana var. elongata A. Braun); Laches, 

2800 m, Lindig s.n. (BM); Sebastopol, Cordillero Oriental 2400 m, Little 8611 (COL); falls of Rio 

Bogota, vicinity of El Salto, 2430-2450 m, Little 7897 (COL). Norte de Santander: La Mesita, 

Pamplona, 2825 m, Alston 7236 (BM). 

ECUADOR. Along Rio Valladolid, Quebrada Honda and Tambo Valladolid, 2-3000 m, Steyermark 

54584 (BM). 

Geographical range: Venezuela, Colombia and Ecuador, at high elevations. 

38. Selaginella flacca Alston, sp. nov. (Figs 7 & 8) 

Species heterophylla ex affinitate S. moritzianae sed difTert caulibus ubique rhizophoros 
gerentibus; habitu laxiore et caulibus flagelliferis. 

Planta ubique repens caulibus glabris, circa 9 cm longis, filiformibus, basi 03 mm in 
diam., e basi ramosis, in siccitate albido-stramineis, foliis ad basin heteromorphis, 
rhizophora ubique gerentibus; parte frondosa ambitu oblonga, ramis circa 0*5 cm inter se 
distantibus, pinnatis, ambitu irregularibus, usque ad 3 cm longis, 1*5 cm latis; Folia lateralia 
patentia, distantia, ovato-elliptica, 2 mm longa, 1*5 mm lata, minute cuspidata; semi-facie 
superiore semi-ovato-elliptica, basi rotundata, minute ciliato-denticulata, alibi distanter 
serrata; semifacie inferiore semi-oblongo-elliptica, basi rotundata, distanter et minute 
serrata; folia axillaria lateralibus similia sed minus obliqua; folia intermedia elliptica, vel 
ovato-elliptica, 1 mm longa, 0*7 mm lata, distantia, distanter serrata, apice acuminato. 
Strobili in apicibus ramulis lateralibus singuli dispositi, platystachyi, 5 mm longi, 2*25 mm 



270 



A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 





- 

■** * 








; -if. 



r 



!*^|p yf 



/7>r 



jf l W..)t. U^w^.xtic 



/v^ 



V * " 









1 vAg* e 



: ft 

felagwtNa -floicca- Alston >W-~' ,$Z 






m^-W 



TYPE SPECIMEN 

SEUGDffiLU FIACCA Alatc 



Fig. 7 Selaginella flacca Alston: Type specimen, £Vm7 1 765 (BM). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



271 




Fig. 8 Selaginella JJacca Alston: A. Close-up of lateral (/) and axillary (a) leaves, x 18. B. Lower 
epidermis of lateral leaf showing stomata and papillate ridges, x 190. C. Upper epidermis of 
lateral leaf showing stoma (s) on margin, x 340. D. Close-up of median leaf, x 1 15. All from 
Ernst 1765. 



lati, non ramosi; sporophylla dimorpha, dorsalia viridia oblique anguste oblongo-lanceolata, 
distanter serrata, apice acuto; ventralia hyalina, aequilateralia anguste oblongo-lanceolata, 
distanter serrata, apice acuminato; megasporae c. 325 /im, luteae, laeves; microsporae c. 
28 //m, luteae (humectae), rugulosae, parce granulosae. 



272 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Typus: Venezuela, Miranda: Quebrada Seburan, prope Caracas, Ernst 1765 (BM, 
holotype). 

Other specimens seen: 

VENEZUELA. Miranda: Galipan, Moritz 71 & 221 (BM); Caracas, Moritz 71 (BM); Sierra de El 

Avila, 1675 m, Steyermark 55135 (BM, US); Silla de Caracas, Ernst 1567a (BM). Sucre : Helvetia, 

1 730 m, at the mouth of Rio Amana, around Turumiquire, Nett 1 1 (BM). 

COLOMBIA. Cundinamarca: Cordillera Oriental, canyon at NE edge of Bogota (E of Calle 72), 

2700 m, Little 9802 (COL). 

Geographical range: The Andean chain of Venezuela and Colombia south to 4°N. 

Notes: It appears that the determiner [A. Braun ?] of Moritz collections sorted them into 
species groups when gatherings were mixed. We therefore find labels bearing 4 71 & 22 T, 
meaning that the material thus distributed with that label came from both field collections. 

39. Selaginella macilenta Baker in /. Bot., Lond. 22 : 90 (1884); Fern Allies: 68 (1887). 
Type from Ecuador, Chimborazo: at the foot of Mt Chimborazo, 900 m, Spruce s.n. (K). 

Specimens seen: 

COLOMBIA. Santander: Carcoba, near Bucaramanga, 2450 m, Alston 7293 (BM). 

ECUADOR. Chimborazo: type as above. Pinchincha: by Rio Toachi, above the confluence with the 

Rio Pilaton, 900 m, Bell 221 (BM); Saloya, W side of the Cordillera Occidental, 1800 m, Acosta Solis 

5790 (BM). 

Geographical range: Colombia and Ecuador. 

Notes: The type specimen has been mounted with the stems matted together so that the form 
of the individual plants is difficult to make out, but the species appears to differ from S. 
cavifolia (No. 37), S. moritziana (No. 42), and S. popayanensis (No. 36) by distinctly aristate 
median leaves, numerous soboles or terminal flagellae, and narrow sporophylls. 

40. Selaginella mollis A. Braun in Annls Sci. nat. (Bot.) V, 3:276 (1865); Baker, Fern 
Allies : 85 (1887). Type from Colombia, Norte de Santander: Ocana, 1800 m, Schlim 1029 
(K). 

Specimens seen: 

COLOMBIA. Antioquia: Rio Mulatos, E of Turbo, 75 m, Haught 4957 (BM, COL). Choco: between 
Atrato and Truando, Schott 8 (BM); between Truando and Nercua, Schott 8 (NY). Norte de Santander: 
type as above. 

Geographical range: Mexico to Colombia. 

Notes: This species resembles S. heterodonta (Desv.) Hieron. of the Greater Antilles, but in 
that species the median leaves are elliptic-oblong with long cilia, while in S. mollis they are 
nearly round and with short cilia. 

41. Selaginella flagellata Spring in Bull. Acad. r. Belg. 10 : 228 (1843); in Mem. Acad. r. Sci. 
Lett. Belg. 24 : 207 (1850); Baker, Fern Allies : 73 (1887); Alston in J. Bot., Lond. 72 : 37 
(1934), in Pulle, Fl. Suriname 1 : 168 (1938). Type from French Guiana, Inini: source of 
the Rio Oyapok, Leprieur s.n. (LG, holotype; P, isotype). 

Selaginella ambigua A. Braun in Annls Sci. nat. (Bot.) V, 3 : 286 (1865); Baker, Fern Allies : 

121 (1887). Type from Venezuela. 
Selaginella leptostachya A. Braun in Annls Sci. nat. (Bot.) V, 3 : 289 (1865); Baker, Fern 

Allies : 122 (1887). Type from Colombia. 
Selaginella regularis Baker in J. Bot., Lond. 22 : 277 (1884); Fern Allies : 78 (1887). Type 

from Peru. 
Selaginella rhizophora Baker in J. Bot., Lond. 22:244 (1884), Fern Allies : 73 (1887). 

Syntypes from Panama. 
Selaginella purdiei Hieron. in Hedwigia 58 : 322 (1917). Type from Trinidad. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 273 

Selaginella othmeri Hieron. in Hedwigia 58 : 324 (1917). Type from Trinidad. 
Selaginella reptans Sodiro, Crypt. Vase. Quit. : 597 (1893). Type from Ecuador. 
Selaginella polysperma sensu Sodiro, Rescens. Crypt. Vase. Quitensis.: 92 (1883), non 
Spring. 

Specimens seen: 

GUYANA. Berbice: Sierra Acarai Region, 1/4 mile south of Camp 3, Guppy 378 (BM); Manicole bog, 
2 miles NE of Camp 1 , Guppy 336A (BM). Essequibo: Kaietur, Jenman ff. (NY). 
FRENCH GUIANA. Inini: type as above. 
SURINAM. Saramacca: Tafelberg, Maguire 24337 (BM). 

VENEZUELA. Aragua: Colonia Tovar, Fendler 324 p.p. (B; syntype of S. ambigua A. Braun); north of 
Maracay, 900 m, Box 3922 (BM); Guamitas, 760 m, Williams & Alston 205 (BM), Alston 5761 (BM). 
Bolivar: near Salto de Pacairao, NE of Santa Teresita de Kavanayen. Steyermark 60523 (BM). 
Carabobo: between Carabobo and Tinaquillo, 500 m, Alston 5747 (BM); above Hacienda Cura, 
1000 m, Alston 6291 (BM). Delta Amacuro: Imataca Mountains, Thompson 20 (BM). Merida: Santa 
Cruz de Zamora, 1000 m, Alston 6992 (BM). Nueva Esparta: Margarita Island, Pico Blanco, 500 m, 
Gines 4074 (BM). Sucre: Quebrada del Imposible, Cumana, Moritz 221 (B, syntype of S. ambigua A. 
Braun). Yaracuy: Hacienda Iboa, near Guyama, 4-500 m, Pittier 1 1 145 (NY, VEN). 
COLOMBIA. Boyaca: Beside trail from Bocota to Bachira, on steep north bank of the Rio Royata, c. 
2050 m, Grubb & Guymer P 95 (BM); Muzo Minas Mujo, 700 m, Lindig 1514 (BM, type-collection of 
S. leptostachya). Magdalena: near Onaca, 750 m, Smith 2245 (BM, NY, US). Meta: Guapayita, 
Macarena, Idrobo & Schultes 833a (BM); Rio Guatriquia near Villa Vicencio, 500 m, Alston 7603 
(BM). Santander: near Barranca Bermeja, 100-500 m, H aught 1970 (BM), 1979 (BM); near Puerto 
Berrio, 100-700 m, Haught 1870 (BM). Valle del Cauca: Rio Engano, Digua Valley, 700 m, Alston 
7878 (BM). 

ECUADOR. Manabi: R Toachi, Sodiro s.n. (P, syntype of S. reptans Sodiro) San Miguel, Sodiro s.n. 
(P, syntype of S. reptans Sodiro). Pichincha: Andes of Quito, Sodiro s.n. (P); near Quinide, Holdridge 
1665 (BM). 

PERU. Cuzco: Sapansachayoc, Paucartambo, 900 m, Vargas 7360 (BM); near Cuzco, Jay a (NY); 
Quellanu, 900 m, Bues s.n. (B). Huanuco: Cueva de las Pavas, 5-600 m, Ferreyra 2892 (BM). Junin: 
Chanchamayo Valley, Schunke 189 (BM); E of Quimin Bridge, La Merced, 700-1300 m, Killip & 
Smith 23945 (BM, NY), 24021 (BM, NY). Loreto: Tarapoto, Spruce 3977 (BM, GCE, type collection 
of S. regularis Baker); between Yurimaguas and Balsapuerto 135-150 m Killip & Smith 28350 (BM, 
NY); above Pongo de Manseriche, 220 m, Mexia 6366a (BM). San Martin: Juan Jui, Alto Rio 
Huallaga, 4-800 m, Klug 4230 (BM, UC). 

BOLIVIA. Cochabamba: Antathuacana, Espiritu Santo, Buchtien 2252 (BM); s.n.(Rosenstock exsicc. 
98; BM). La Paz: Guanay, Tate 557 (NY); Sorata, Larecaja, 300, Rusbv 203 (NY); Yungas, 1800 m, 
Rusby 453 (NY). 

BRAZIL. Amazonas: Bocco de Tejo, Rio Jurua, Vie 5524 (K). Matto Grosso: Poaia, Buritzinho, 
Lindman A 3265 (NY). 

Geographical range: From Mexico to Bolivia and east to French Guiana and Brazil. 

Notes: Braun states that the microspores of his S. leptostachya are tuberculate (not papillose); 
this is true but they appear to be unripe. This species may be separated from S. cladorrhizans 
(No. 43) by its more ovate, acute, cordate leaves. S. muscosa (No. 88) has similar leaves, but 
the strobili are tetragonous; its habit is prostrate and it lacks flagellae. S. pearcei (No. 34) is 
darker green with shortly aristate median leaves, S. flagellata being more delicate, paler 
green with long aristate median leaves. S. sandwithii (No. 80) is more erect, lacks flagellae 
and has tuberculate microspores, whilst those of S. flagellata bear elongate papillae as in S. 
cladorrhizans. 

42. Selaginella moritziana Spring ex Klotzsch in Linnaea 20 : 436 (1847), nom. nud., in 
Mem. Acad. r. Sci. Lett. Belg. 24 : 249 (1850); Baker, Fern Allies : 69 (1887), p.p. quoad pi. 
venezuelensis. Type from Venezuela, Merida: without exact locality, Moritz 311 (BM). 

Selaginella moritziana var. normalis A. Braun in Annls Sci. nat. (Bot.) V, 3 : 284 (1865). 
Type as above. 



274 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Specimens seen: 

VENEZUELA. Federal District: Los Flores, Serra de El Avila, 1600 m, Alston 5538 (BM). Merida: 

type as above. 

ECUADOR. Pichincha/Napo-Pastaza: vallee de Lloa (Ungui), Benoist 2700 (BM). 

Geographical range: Northern Venezuela and Ecuador. 

Notes: Braun (1865: 283) (under S. cladorrhizans) writes of Fendler 324 'mixta cum S. 
ambigua et moritziana\ S. cavifolia (No. 37) from the Tequendama Falls near Bogota, has 
more obtuse, relatively broader lateral leaves, scarcely acuminate median leaves, and 
broader sporophylls. S. macilenta (No. 39), from 900 m at the base of Chimborazo, appears 
to differ by narrower, more oblong lateral leaves, and aristate median leaves. S. potaroensis 
(No. 87) has relatively smaller, much tapered, more ovate, unequal-sided median leaves. S. 
flacca (No. 38) has more distant, paler, green lateral leaves, and the stem bears rhizophores 
throughout. Spring erroneously cited the type as from Colombia. 

43. Selaginella cladorrhizans A. Braun in Annls Sci. nat. (Bot.) V, 3 : 282 (1865); Baker, Fern 
Allies : 72 (1887); Alston in J. Bot., Lond. 72 : 37 (1934). Types from Venezuela, Aragua: 
Colonia Tovar, Fendler 324 p.p. (B, BM) and Falcon: San Carlos, Moritz 448 (B,BM). 

Specimens seen: 

FRENCH GUIANA. Podidier: Leprieur 166 (K). 

VENEZUELA. Anzoategui: confluence of Rio Leon and Rio Zumbador, 4-500 m, Steyermark 61191 

(BM). Aragua: Colonia Tovar, Moritz 71? (B): syntype, as above. Carabobo: between Carabobo 

Tinaquillo, 500 m, Alston 5746 (BM); Hacienda Stelling, 700 m, Alston 6300 (BM). Delta Amacuro: 

Imataca Mts, Thompson 19 (BM). Falcon: syntype as above. Miranda: Galipan, Moritz s.n. (BM); 

Petare, Williams & Alston 2 1 3 (BM); between Petare and La Guayrita, 900 m, Alston 5469 (BM). 

COLOMBIA. Magdalena: Minea, Santa Marta, 600 m, Smith 2567 (NY, US). Vaupes: Soratama, 

between Rio Pacoa and Rio Kananari, Schultes & Cabrera 12755 (COL, US). 

BRAZIL. Para: Limestone rocks at Trovadore, Traill 1436 (K). 

Geographical range: Mexico southwards to French Guiana and Brazil, and in Colombia; also 
recorded for Trinidad. 

Notes: A specimen at New York labelled Guiana, Jenman, was more probably from 
Trinidad. It may be necessary to reduce this species to S. tenella (Beauv.) Spring of the 
Antilles. It is similar in habit to S. lychnuchus (No. 47), but that species has strongly 
dimorphous sporophylls. 

44. Selaginella porphyrospora A. Braun in Annls Sci. nat. (Bot.) V, 3 : 286 (1865). Type 
from Mexico: Vera Cruz, Sartorius s.n. (B, BM). 

Selaginella bulbifera Baker in Gdnrs' Chron. 1867 : 783, 950 (1867). Type from garden 

material of unknown origin. 
Selaginella binervis Liebm. ex Fourn., Mex. PL : 148 (1872), in syn. 
Selaginella albonitens sensu Christ in Dur. & Pittier, Primit. FL Costaric. 1 3 : 255 (1896), 

non Spring. 
Selaginella bernoulii Hieron. in Hedwigia 41 : 192 (1902). Type from Guatemala. 

Specimen seen: 

COLOMBIA. Boyaca: beside trail from Bocota to Bachira, on steep north bank of the Rio Royata, c. 

2050 m, Grubb & Guymer P.95 (BM, COL). 

Geographical range: Mexico south to Colombia. 

45. Selaginella novae-hollandiae (Sw.) Spring in Bull. Acad. r. Belg. 10 : 234 (1843), in Mem. 
Acad. r. Sci. Lett. Belg. 24 : 208 ( 1 850). Type from 'Nova Granada'? [see Notes]. 

Lycopodium novae-hollandiae Sw., Syn. Fil. : 1 84, 410(1 806). Type as above. 
Lycopodium ciliatum Willd., Sp. PL 5:38 (1810), nomen abortivum, non Lam. (1778). 
Specimen from l Nova Granada'? [see Notes]. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 275 

Selaginella increscentifolia Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 106 (1850). Syntypes 

from Ecuador, Bolivia and Peru. 
Selaginella warscewicziana Klotzsch [ex Lauche], Verz. August. Garten :8 (1856), nom. 

nud.; Kummerle in Mag. Bot. Lap. 13 : 52 (1915), nom. nud. 
Selaginella warscewiczii Linden, Cat. No. 12 : 23 (1857), nom. nud.; Klotzsch ex A. Braun 

in Annls Sci. nat. (Bot.) V, 3 : 276 (1865). 
Selaginella ciliata (Willd.) A. Braun, Index Sem. Hort. bot., Berl. Appendix 1857 : 16(1857), 

non Opiz (1823). Type as above. 
Selaginella radiata sensu Baker, Fern Allies : 86 (1887); Sodiro, Crypt. Vase. Quit. : 614 

(1892), non Spring (1843). 
Selaginella triuncialis Sodiro, Crypt. Vase. Quit. : 594 (1893). Type from Ecuador. 
Selaginella tucumanensis Hieron. in Bot. Jb. 22 : 419 (1896). Type from Argentina. 

Specimens seen: 

VENEZUELA. Carabobo: La Mona, near Chirgua, 700 m, Alston 5940 (BM). Merida: between 
Hacienda Agua Blanca, above La Azulita, and Rio Capaz, 975 m, Steyermark 56 1 50 (BM). 
COLOMBIA. Antioquia: Amaya, Daniel 4250 (BM); near Antioquia, 550 m, Barclay & Arboleda 9 
(BM). Choco: Trunado Fails, Schott 1 1 (BM). Cundinamarca: between Pacho and La Palma, 1 100 m, 
Haught 6010 (BM, COL); between Villeta and Guaduas, 1000 m, Haught 6744 (BM, COL); near 
Bogota, 1000-1400 m, Triana s.n. (BM); Carretera between Santandercito and El Salto, 2200 m, 
Pinto, Polidoro & Dumont 585 (COL). Meta: Villavicencia, 500 m, Alston 7613 (BM); Rio Guapaya, 
475 m, Philipson, Idrobo & Fernandez 1678 (BM); Guapayita, Macarena 5-600 m Idrobo & Schultes 
833 (BM). Valle del Cauca: Cali, Andre 441 (NY); Cisneros, Dagua Valley, 300-500 m Killip 1 1497 
(BM, NY), 35601 (COL); Dagua 700-900 m, Killip 5549 (BM, NY); 200-300 m, Lehmann 8913 (NY); 
Rio Engano, Dagua valley, 700 m, Alston 7818 (BM), 7879 (BM); Rio Dagua valley, Chorrera, La Elsa, 
975 m, Killip 34802 (COL); La Margarita, c. 760 m, Killip 34903 (COL). 

ECUADOR. Without precise locality: Fraser s.n. (BM); Lindig 1517 (BM); Andes of Quito, Couthouy 
83 (NY); Jameson 13 (BM); Nibey (Mbey ?), Sodiro s.n. (NY); way to Nanegal, 2200 m, Mille s.n. 
(NY);Niebli,^dr<?3771 (NY); San Florencio, 1580 m, Andre 3771 (NY); San Jose de Toachi, 100 km 
W of Quito, c. 1000 m, Bell 190 (BM). Azuay: W of Patal, between Huahualcay and Passas de Pinglion, 
2670-3275 m, Steyermark 52601 (BM). Bolivar: San Christobal, 1070 m, Asplund 8348 (S). 
Chimborazo: Near Huigra, R. Chanchan, 1350 m, Camp E3131 (BM); 1500-1950 m, Camp E 3275 
(BM). Pichincha: Jameson s.n. (K); Canzacoto, Sodiro s.n. (P, ? type, labelled S. triuncialis Sod.); Santo 
Domingo de los Colorados, 900 m, Holdridge 1589 (BM); Chaupi-Sagcha, Puluagua, c. 1800 m, Bell 
473 (BM); Nono, 2600 m, Asplund 7481 (S). Santiago-Zamora: Above Rio Upano, 585-660 m, Camp 
E 1448 (BM); Macas, Rio Upano, Asplund 19817 (S); near Mendez, 570 m, Camp E 894 (BM); E 101 1 
(BM); E 1475 (BM); E 1476 (BM). Tungurahua: Bell 828 (BM); between Banos and Rio Verde, c. 
1650 m, Agoyan, Asplund 7598 (S). On border between Loja and Santiago-Zamora: crest of the 
Cordillera de Zamora, east of Loja, c. 3000 m, Camp E 8 1 (BM). 

PERU. Amazonas: Chachapoyas, Mathews s.n. (BM). Ayacucho: Pampalca, between Huanta and Rio 
Apurimac, 3200 m, Killip & Smith 22227 (BM, NY), 23240 (BM, NY); Aina, between Huanta and Rio 
Apurimac, 750-1000 m, Killip & Smith 23177 (BM, NY), 22545 (BM, NY); Coarrapa, between 
Huanta and Rio Apurimac, 1500 m, Killip & Smith 22364 (BM, NY). Cuzco: Machupicchu, 2400 m, 
Herrera 3475 (BM); Stafford 1068 (BM); Torontoy, Urubamba Valley, 2400 m, Cook & Gilbert 1 190 
(US); San Miguel, Urubamba Valley, 1800 m, Cook & Gilbert 1004 (US); 1 169 (US). Junin: Mathews 
1084 (BM, NY, US); Huacapistana, 1800 m, Killip & Smith 24275 (BM, NY); 10 km SW of San 
Ramon, 1000 m, Trvon 5448 (BM). La Libertad: Otuzco, road to Paranday (Sinsicap), 2600 m, Lopez 
1044 (BM). Loreto: Balsapuerto, lower R. Huallaga, 350-550 m, Killip & Smith 28488 (BM, NY); near 
Tarapato, Spruce 2328 (BM, CGE). Piura: near Ayabaca, 2700 m, Berry s.n. (US). Puno: Ollachea, 
prov.Carabayo,2500 m, Vargas 69 \4(BM). 

BOLIVIA. Without precise locality: Bang 35 (BM); Bridges s.n. (BM). Beni: Huachi, head of Rio Beni 
540 m, White 541 (BM, NY, US). Cochambamba: Lagnillas, over mountains behind Choro in St Elena 
valley, 3000 m, Brooke 6256 (BM); Incachaca (small power station) c. 80 mis NE of Cochamba, 
2400 m, Brooke 6673 (BM). La Paz: near La Paz, 3000 m, Rusby 453 (NY, US); 3600 m, Rusby s.n. 
(BM); Queliquaya, Sorata, Larecaja, 2700 m, Mandon 1530 (BM, NY); Hacienda Simaco on way to 
Tipuam, 1400 m, Buchtien 5277 (US), 5278 p.p. (US), 5279 (US); Yungas, Bang 440 (BM, NY); 
1200 m, Rusbv 203 (NY); Paradiso, 1350 m, Williams 1402 (NY, US). Santa Cruz: Samaipata, 
1800 m, Steinbach 3740 (BM). Tarija: Pinos, near Tarija 2300 m, Fiebrig2918 (BM). 



276 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Geographical range: Nicaragua southwards to Bolivia. 

Notes: S. novae- hollandiae is one of the commonest species of Selaginella in South America. 
It is very variable in the density and length of cilia on the leaves. S. brevifolia (No. 79) is 
distinguished by its more prostrate habit, with rhizophores above the middle, and a 
prominent central ridge on the upper surface of the stem in dried specimens; in S. novae- 
hollandiae there are usually two ridges which are less distinct; furthermore the median leaves 
of S. brevifolia are exauriculate. S. leucoloma (No. 58) is similar in habit, but the median 
leaves are acute (not aristate), exauriculate and more conspicuously white-margined. S. 
popayanensis (No. 36) is separable by its dentate leaves; those of S. novae- hollandiae have 
some long cilia at the base, and are concave below when dry, being strongly cordate and 
overlapping the stem. S. radiata (No. 29), with which it has been confused in the past, is a 
larger plant with more rounded long-aristate median leaves, and lateral leaves which are 
more silvery beneath. The type specimens of Lycopodium novae- hollandiae Sw. and L. 
ciliatum Willd. were both localised 'Nova Hollandia', perhaps a mistake for Nova Granada 
(Colombia), as Australia is certainly not implied. There is a specimen at Paris from 
Canzacito, labelled S. triuncialis Sodiro, but the type locality was on the slopes of Mt 
Atacatzo, Ecuador. It seems to be a small form of S. novae- hollandiae. 

46. Selaginella chionoloma Alston ex Crabbe & Jermy in Am. Fern J. 63 : 137 (1973). 
Type from Peru, Cuzco: Valle de Lares, Calca Cuquipata, Herrera 1636 (US, holotype; 
BM, isotype). 

Specimens seen: 

COLOMBIA. Caqueta: Florencia, Arbelaez 627 (BM, US). 

PERU. Cuzco: type, as above. San Martin: beyond Las Palmas, near tunnel, Tingo Maria, 800 m, 

Allard 21 130 (BM). 

BOLIVIA. La Paz: Yanacachi, Yungas, Jay b (US). 

Geographical range: Colombia and Peru to north-western Bolivia. 

Notes: This species resembles S. ramosissima (No. 48) in general appearance, but that 
species has a stouter stem and lacks soboles and the conspicuous white margin of the lateral 
leaves, which is a feature of S. chionoloma. It is most closely related to S. novae- hollandiae 
(No. 45), differing only in its lack of definite cilia on the lateral leaves, its more pronounced 
white border, especially on the acroscopic edge of those leaves, and its narrower median 
leaves. Pierre Jay's specimen is very poor, but seems to be this species. 

47. Selaginella lychnuchus Spring [ex Klotzsch in Linnaea 20 : 435 (1847), nom. nud.], in 
Mem. Acad. r. Sci. Lett. Belg. 24 : 247 (1850); Baker, Fern Allies : 121 (1887). Syntypes 
from Venezuela, Federal Dist.: Galipan, Moritz 71 2 (B, BM, LG); Merida, Moritz 378 (B, 
BM, LG). 

Selaginella lychnuchus var. flaccida Spring [ex Klotzsch in Linnaea 20 : 436 (1847), nom. 

nud.], in Mem. Acad. r. Sci. Lett. Belg. 24 : 248 (1850). Type as above (Moritz 71 2 ). 
Selaginella lychnuchus var. rigidiuscula Spring [ex Klotzsch in Linnaea 20:436 (1847), 

nom. nud.], in Mem. Acad. r. Sci. Lett. Belg. 24 : 248 (1850). Type as above (Moritz 378). 
Selaginella lychnuchus var. pusilla A. Braun in Annls Sci. Nat. (Bot.) V, 3 : 288 (1865). Type 

from Venezuela. 

Specimens seen: 

VENEZUELA. Aragua: below Rancho Grande, 1000 m, Williams & Alston 144 (BM); near La 

Regriesiva, km 25, 1000 m, Williams & Alston 170 (BM). Federal District: syntype, as above. Merida: 

syntypes as above; Rio Albariga, 2400 m, Alston 6969 (BM). Sucre: Quebrada del Impossible, Moritz 

221 &71 (BM). 

COLOMBIA. Santander: Corcoba near Bucaramanga, 2200 m, Alston 73 1 1 (BM). 

Geographical range: Costa Rica to Colombia and Venezuela. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 277 

Notes: There is some confusion over Moritz's field numbers, and the reason for double 
numbers is not clear; but there is no ambiguity. There is no doubt that the gatherings were 
mixed [see S. cladorrhizans (No. 43), S. jlacca (No. 38), S. flagellata (No. 41) and S. 
pallescens (No. 5)], and the double numbers may have arisen when Spring or Moritz (or A. 
Braun?; see p. 272) were sorting them out. On original labels (at least at BM) the two 
numbers are linked by an ampersand (&). 

Three specimens of Moritz are cited here, although Spring (I.e.) only cites two and those as 
varieties which are solely habitat forms and extremes of the morphological range. Spring 
gives Colombia as the locality, in error. 

48. Selaginella ramosissima Baker in J. Bot., Lond. 23 : 295 (1885); Baker, Fern Allies : 12 
(1887); Knox in Trans, bot. Soc. Edinb. 35 : 260 (1950). Type from Peru, Loreto: near 
Tarapoto, Spruce 4088 (K, holotype; BM, CGE, NY, isotypes). 

Specimens seen: 

ECUADOR. Santiago-Zamora: between Rio Sabamba and Canillones Tambo, 1300-2100 m, Camp 

E66(BM). 

PERU. Cajamarca: Monte Seco, Hualgayoc, 1800 m, Soukup 3872 (BM). Loreto: type as above. 

Geographical range: Ecuador to Peru. 

49. Selaginella glossophylla Alston ex Crabbe & Jermy in Fern Gaz. 11 : 259 (1976). Type 
from Bolivia, Cochabamba: Incachaca, 80 miles NE of Cochabamba, 3000 m, Brooke 
6783 (BM). 

Specimens seen: 

CHILE. Without exact locality: Mercier s.n. (P). 

BOLIVIA: type as above. 

Geographical range: Bolivia and Chile. 

50. Selaginella minima Spring in Bull. Acad. r. Belg. 10 : 139 (1843); Mem. Acad. r. Sci. Lett. 
Belg. 24 : 86 (1850); Baker, Fern Allies : 84 (1887). Type from French Guiana: ad basin 
montis Eigre Insulae Cayenne, Leprieur 1 58 (G, K, P, US). Only specimen seen. 

Geographical range: Costa Rica, Panama, French Guiana. 

Notes: This species is somewhat doubtfully distinct from S. broadwayi Hieron., described 
from Trinidad, and S. simplex (No. 51). In S. broadwayi the lateral leaves are not ciliate, but 
otherwise similar; in S. simplex the lateral leaves are ovate and denticulate. The three 
species may form a cline. 

51. Selaginella simplex Baker, in J. Bot., Lond. 23 : 293 (1885), Fern Allies : 121 (1887). 
Type from Brazil, Para: near Santarem, Spruce 947 (BM, CGE, K). 

Specimens seen: 

VENEZUELA. Carabobo: La Mona, near Chirgua, 700 m, Alston 5959 (BM). 

BOLIVIA. Near Santa Cruz, Williams 2657 (BM, NY, US). Goias: Cerrado 8 km S of Cristalina 

1200 m, Irwin, Grear, Souza & Reis dos Santos 1 3654 (BM). 

BRAZIL. Para: Type as above. Ceara: Cedro, Loefgren 1 160 (BM); Pernambuco: Tapera, wall of ditch, 

Picket 281 2 (BM). 

Geographical range: Trinidad, Venezuela, Bolivia, Brazil. 

52. Selaginella tyleri A. C. Smith in Bull. Torrey bot. Club 58 : 31 1 (1931). Type from 
Venezuela, Bolivar: slopes of ridge 24, Mount Duida, 1680 m, Tate 463 (BM [fragment], 
NY). Only specimen seen. 

Geographical range: Confined to Mt Duida. 



278 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

53. Selaginella cruciformis Alston ex Crabbe & Jermy in Fern Gaz. 11 : 257 (1976). Type 
from Venezuela, Carabobo: Rio Aguada, 1500 m, Alston 64240 (BM, holotype; AAU, 
COL, INPA, QCA, RB, SP, U, USM, US, VEN, isotypes). 

Specimens seen: 

VENEZUELA. Aragua: Rancho Grande, 1400 m, Box 3879 (BM); Tschudi 64 (US). Carabobo: type as 
above. Merida: prope Tovar, 900 m, Fendler 487 (BM). Yaracuy: Cerro La chapa, north of Nirgua, 
1200-1400 m, Steyermark, Bunting & Wessels-Boer 100259 (BM, US). 

Geographical range: Confined to Venezuela. 

Notes: This species resembles S. chrysoleuca (No. 55) in the sessile lateral strobili and the 
basal rhizophores. It might also be confused with S. truncata (No. 94), but that species is 
prostrate, rooting throughout, and has terminal strobili. 

54. Selaginella acanthostachys Baker in J. Bot., Lond. 21 : 99 (1883), Fern Allies : 44 (1887). 
Type from Peru, Loreto: Monte Campana, Spruce 4328 (K). 

Specimens seen: 

ECUADOR. Santiago-Zamora: Valley of R. Negro, down to R. Pailas, Camp E 4952 (BM); Cordillera 

de Zamora, Loja, 3000 m, Camp E 80 (BM). 

PERU. Cajamarca: Tambillo, Cutervo, 2190 m,Raimondi 679\ (BM). Loreto: Type as above. 

Geographical range: Ecuador and Peru. 

55. Selaginella chrysoleuca Spring in Bull. Acad. r. Belg. 10 : 226 (1843), in Mem. Acad. r. 
Sci. Lett. Belg. 24 : 197 (1850); A. Braun in Annals Sci. nat. (Bot.) V, 3 : 274 (1865); Baker, 
Fern Allies : 81 (1887). Type from Bolivia, without precise locality, D'Orbigny s.n. (P). 

Selaginella sprucei Hook., Second Cent. Ferns t.83 ( 1 86 1 ). Type from Peru. 

Specimens seen: 

VENEZUELA. Aragua: near Puerto Cabello, Karsten s.n. (B). 

COLOMBIA. Santander: Mesa de los Santos, 1500 m, Killip & Smith 15211 (B, BM, NY, US); 

Coromoro, ENE of Charala, 2300 m, Ewan 1 5664 (BM). 

ECUADOR. Santiago-Zamora: between Rio Ontza and Chupiasa, Cordillera Cutucu, 1250-1300 m, 

CampE 1 196 (BM); SW of Rio Itzintza, Cordillera Cutucu, 1500-1600 m, Camp E 1388 (BM). 

PERU. Junin: Pichis trail, Porvenir, 1500-1900 m, Killip & Smith 25951 (NY, US). Loreto: Mt 

Campana, near Tarapoto, Spruce 4623 (BM, CGE, K; type of S. sprucei); between Tingo Maria and 

Pucallpa, Ferreyra 1008 (BM). 

BOLIVIA. Without precise locality: type as above. La Paz: Lower Rio Pelichuco, Caupolican, 1200 m, 

Williams 2647 (NY, US); Santa Barbara, Caupolican, 1650 m, Williams 1395 (NY, US); Hacienda 

Simaco, on way to Tipuani, Larecaja, 1400 m, Buchtien 527 '4 (BM, NY, US); Polo-Polo near Coroica, 

N Yungas, 1 100 m, Buchtien 3549 (US). 

Geographical range: Venezuela to Boliva. 

Notes: The habit of this species resembles that of S. cruciformis (No. 53); but the median 
leaves on the former are falcate with apices crossing one another, and those of the latter 
subacute with the apices directed forwards. The specimens from Colombia and Venezuela 
are smaller and perhaps represent a distinct species. 

56. Selaginella contigua Baker in J. Bot., Lond. 22:295 (1884), Fern Allies: 79 (1887); 
Hieron. in Hedwigia 43 : 44 (1903). Syntypes from Brazil, Rio de Janeiro, Glaziou 4493 
(C, K) and Itatiaia, Glaziou 5638 (C, K). 

Selaginella callimorpha Silveira in Bolm Comm. geogr. geol. Minas Geraes 5:121 

(1898). Type from Brazil. 
Selaginella wettsteinii Hieron. in Denkschr. Akad. Wiss. Wien 79 : 60 (1908). Type from 

Brazil. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 279 

Specimens seen: 

BRAZIL. Without exact locality: Glaziou 52 1 6 (C), 56 1 3 (P); Porta d'Estrella el Mandioca, Beyrich s.n. 
(BM). Minas Gerais: Rio Novo, Capanema s.n. (BM); Serra de Bocaivra, Schwacke s.n. (BM). Parana: 
Schwacke s.n. (BM). Rio de Janeiro: syntypes as above; Glaziou 9312 (C, K, P), 9314 (C, K); Mosen 53 
(K); Tyuca, Hoehne 24876 (P, S); Glaziou 4676 (C, P); Lutz 1151 (BM); Alto da Boa Vista de Nova 
Friburgo, Glaziou 7283 (BM); Mt Itatiaya, 800 m, Smith 1638 (C); Itatiaia, Rose & Russell 20600 
(NY); Itatiaia, Angra dos Rios, Maromba, Brade 14530 (BM), 14531 (BM); Serra do Itatiaia, 100 m, 
Brade 6506 (BM); Dusen 708 (BM, P); Sampaio 4052 (BM); Represa Camorima, Brade 12564 (BM); 
Santa Catarina : Joinville, Muller 133a (NY). Sao Paulo: Serra do Cubata, Guillemin 578 (P); Santos, 
Mosen 3556 (K), 3814 (C); Edwall 164 (BM); Rose & Russell 2\ 128 (NY); ?Jap-nyba, Hoehne & Gehrt 
s.n. (BM); Alto da Serra, Wacket 20997 (BM, P); Campo Grande, Sera do Mar, Brade 6600 (NY); S. 
Jose de Barreiro, Loefgren & Edwall 20991 (BM, P); Brasso near Itapecirica, 1000 m, Wettstein & 
Schnifner s.n. (W, type of S. wettsteinii Hieron.); Iguape, 800 m, Brade 8628 (BM); Serra da Bocaina, 
Schwacke & Glaziou s.n. (BM); Itatinga, Brade 10047 (BM), 10264 (BM). 

Geographical range: Confined to south-eastern Brazil. 

Notes: S. callimorpha A. Silveira was reduced to synonymy by Hieronymus in Hedwigia, 
43 : 44 (1904); we have not seen the type. 

57. Selaginella mendoncae Hieron. in Engl. & Prant, Nat. Pflanzenf. 1 (4) : 693 (1901), in 
Hedwigia 45 : 43 (1904). Type from Brazil, Rio de Janeiro, Mendonca 302 (K). 

Specimens seen: 

BRAZIL. Without exact locality: Glaziou 7283 (C). Rio de Janeiro: type as above; Glaziou 12294 (BM, 

C,K). 

Geographical range: Apparently confined to the area around Rio de Janeiro. 

58. Selaginella leucoloma Alston ex Crabbe & Jermy in Fern Gaz. 11 : 262 (1976). Type 
from Bolivia, La Paz: Hacienda Simaco near Tipuani, Larecaja, 1400 m, Buchtein 5280 
(BM, holotype; US). 

Specimens seen: 

BOLIVIA. La Paz: Peluchuco Weddell 4779 (P); type as above and same locality, Buchtein 5278A 

(BM,US). 

Geographical range: Endemic to Bolivia. 

Notes: In habit resembles S. popayanensis (No. 36), but has ciliate, obtuse, lateral leaves. 

59. Selaginella xiphophylla Baker in J. Bot., Lond. 22 : 296 (1884), Fern Allies : 80 (1887). 
Type from Peru, Loreto: Mt Guayrapurima, near Tarapoto, Spruce 3990 (K, holotype; 
BM, CGE, NY, isotypes). Only specimen seen. 

Geographical range: Confined to Mt Guayrapurima, Peru. 

60. Selaginella substipitata Spring in Bull. Acad. r. Belg. 10 : 227 (1 843); Baker, Fern Allies : 
58(1887). Type from Guadeloupe, Beaupertius s.n. (? BR, not seen). 

Selaginella rigidula Baker in J. Bot, Lond. 22 : 295 (1884), Fern Allies : 79 (1887). Type 

from Colombia. 
Selaginella karsteniana A. Braun in Annls Sci. nat. (Bot.) V, 3 : 288 (1865); Baker, Fern 

Allies : 122 (1887). Type from Venezuela. 

Specimens seen: 

VENEZUELA. Without precise locality: Karsten s.n. (BM); Aragua: La Regresiva, 1000 m. Williams 

& Alston 171 (BM). Bolivar: Ptari-Tepui, 2400—2410 m, Steyermark 59604 (BM). Carabobo: Puerto 

Cabello, Karsten 1 74 (B, type of S. karsteniana A. Braun). 

COLOMBIA. Caldas: Rio Santa Rita, Salento, 1600-1800 m, Killip & Hazen 8959 (NY). Santander: 

Ocafia to Pamplona, Kalbreyer912 (K, type of S. rigidula Baker). 

Geographical range: Confined to the cordillera in northern Colombia and Venezuela. 



280 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

61. Selaginella scalariformis A. C. Smith in Bull. Torrey bot. Club 58 : 314 (1931). Type 
from Venezuela, Amazonas: Gorge of Cano Negro, Savanna Hills, Duida, 1200 m, Tate 
8 1 7 (NY, holotype; BM, K, fragment). 

Specimens seen: 

VENEZUELA. Amazonas: type, as above; summit of Cerro Duida, 1025-1200 m, Steyermark 58223 

(BM). Bolivar: N Ridge escarpment of Mt Roraima, 1980 m, Edwards KER61 (BM, K). 

Geographical range: Confined to the sandstone mountains in the south of Venezuela, and 
only recently discovered on Roraima in 1978. 

62. Selaginella wurdackii Alston, sp. nov. (Fig. 9) 

Species heterophylla ex affinitate S. rorairnensis, sed foliis intermediis brevissime aristatis, 
sporophyllis lanceolato-deltoideis, rhizophoris brevibus filiformibus et caulibus brevioribus 
differt. 

Planta ascendens; caulibus glabris, circa 5 cm altis, basi 0*6 mm in diam., parte inferiore 
simplici c. 15 mm; rhizophoris ad quartam partem basin versus restrictis; parte ramosa 
ambitu irregulariter ovato-elliptica, ramis c. 5 mm inter se distantibus, simplicibus vel 
furcatis, usque ad 1 5 mm longis. Folia ubique heteromorpha; folia lateralia patentia, 
distantia oblonga 2 mm longa, 1 mm lata, subacuta; semifacie superiore semi-elliptico- 
oblonga, basi rotundato-cuneata, integra; semifacie inferiore, semi-oblonga, basi cuneata, 
apicem versus distanter serrulata, aliter integra; folia axillaria minus inaequilateralia sed 
aliter lateralibus similia; /o//a intermedia ovato-oblonga, 1 mm longa, 0*75 mm lata, serrata, 
apice breviter aristato. Strobili in apicibus ramulorum, singuli vel bini dispositi, tetragoni, 
4-12 mm longi, 1*8 mm lati non ramosi; sporophylla lanceolata, carinata, distante 
serrulata, apice leviter attenuato, acutoque; megasporae c. 230 //m, luteae (humectae), 
reticulorugosae; microsporae c. 25 jum luteae (humectae) verrucosae. 

Typus: Venezuela, Bolivar: between 4 La Laja' Base Camp, Rio Tirica, Chimanta Massif, 
485-490 m, Steyermark & Wurdack 173 (BM, holotype; US, isotype). 

Other specimens seen: 

VENEZUELA. Bolivar: near Salto de Pacairo, NE of Santa Teresita de Kavanayen, 1220 m, Steyer- 
mark 60499 (BM). 

Geographical range: Known only from south-east Venezuela. 

63. Selaginella seemannii Baker in J. Bot., Lond. 21 : 244 (1883), Fern Allies : 57 (1887); 
Alston in Pulle, Fl. Suriname 1 : 168 (1938). Type from Colombia, Choco: Cacagual Is., 
Seeman 1006 (K, holotype; BM, isotype). 

Selaginella barbacoasensis Hieron. in Hedwigia 43:46 (1904), in Engler, Bot. Jarhb. 
34 : 580 (1 905). Type from Colombia. 

Specimens seen: 

SURINAM. Nickerie: Upper Sipaliwini R., Camp 19, Morro Grande, Rombouts 489 (BM). 

COLOMBIA. Boyaca: Sierra Nevada de Cocuy, in moss forest around Cobugon, c. 2900 m, Grubb & 

Guvmer P 57 (COL). Choco: type as above; Marino: near Barbacoas, Lehmann 89 (BM, type 

collection of S. barbacoasensis Hieron.); Alston 8509 (BM). Norte de Santander : camp 84 on 

pipeline, 540 m, Foster 1722 (COL.). Valle del Cauca: Bridge over Dagua, near Buenaventura, 

Alston 8632 (BM). 

ECUADOR. Napo-Pastaza: near Canelos, 300 — 400 m, Mexia 6906a (BM). Santiago-Zamora; above 

Rio Chupiantza, 585-660 m, Camp E 1444 (BM); above Rio Upano, near Mendez 325-750 m, 

Camp E 972 (BM); Cordillera Cutucu, 1050-1 1 10 m, Camp E 1225 (BM). 

PERU. Loreto: Rio Itaya, c. 10 km S of Iquitos, Tryon 5195 (BM); along R. Itaya, Williams 238 

(BM); Soledad, Rio Itaya, 110 m, Killip & Smith 29638 (NY, US); between Manseriche and mouth of 

the Rio Morona, 1 50 m, Killip, Smith & Dennis 29 \4\ (US). 

Geographical range: Colombia east to Surinam and south to Peru. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



281 




Fig. 9 Selaginella wurdackii Alston: A. Type specimen, Steyermark & Wurdack 173 (BM). B. 
Close-up of lateral (/) and median (m) leaves, x 33. 



282 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Notes: Baker gives the type locality as Panama, but the specimen is from the island of 
Cacagual, which is in Colombia. 

64. Selaginella erectifolia Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 12 (1850); Baker, 
Fern Allies : 75 (1887). Type from Brazil, Rio de Janeiro, Swainson s.n. (K). 

Selaginella camptostachys Fee, Crypt. Bresil : 299 (1869). Type from Brazil. 
Selaginella glazioviana Hieron. in Hedwigia 43 : 36 (1904). Type from Brazil. 

Specimens seen: 

BRAZIL. Minas Gerais: Serra do Itacolumy, Badini 316 (BM). Rio de Janeiro: type as above; Glaziou 
2242 (C, K, type of S. camptostachys Fee); Bocca do Matto (towards Nova Friburgo), 200 — 250 m, 
Lutz 1319 (BM). Santa Catarina; Fachina near Biguassu, Rambo 50378 (BM). 

Geographical range: Confined to southern Brazil. 

65. Selaginella roraimensis Baker in Timehri, 5 : 22 1 (1886), In Trans. Linn. Soc. Lond. 
(Bot.) II, 2 : 295 (1887), Fern Allies : 86 (1887). Type from Venezuela, Bolivar: Roraima, 
im Thurn 122 (BM). 

Specimens seen: 

GUYANA. Essequibo: Macouria Creek, Essequibo River, Jenman 2324 (BM) s.n. (NY). 

VENEZUELA. Bolivar: near Mt Roraima, Jenman s.n. (NY); type as above. 

Geographical range: Confined to the Roraima massif and west Guyana. 

Notes: The specimens from Macouria Creek have slightly broader leaves and a more diffuse 
habit, but do not seem specifically separable. S. porelloides (No. 103) is a smaller species, 
with ovate lateral leaves and median leaves which are aristate or acuminate; S. rhodostachya 
(No. 86) has more ovate lateral leaves and a more creeping habit. 

66. Selaginella scintillata Alston, sp. nov. (Figs 10 & 11) 

Species heterophylla ex affinitate S. roraimensis, sed minus ramosa et foliis minoribus et 
rhizophoris brevibus filiformibus differt. 

Planta ascendens; caulibus glabris, circa 7 cm altis, basi 0*75 mm in diam., e basi ramosis, 
rhizophoris ad tertiam partem basin versus restrictis; parte ramosa ambitu anguste elliptica, 
ramis c. 6 mm inter se distantibus, simplicibus vel irregulariter et distante pinnatis, 
plerumque c. 7 mm longis, sed usque ad 5 cm. Folia ubique heteromorpha; folia lateralia 
plusminusve patentia, contigua, leviter imbricata vel distantia, ovato-oblonga, F5mm 
longa, fere 1 mm lata, subobtusa; semifacie superiore semi-ovata, basi rotundata, crebre 
serrata, apicem versus integra, aliter distante serrulata; semifacie inferiore semi-oblonga, basi 
vix cuneata, integra; folia axillaria lateralibus subsimilia; folia intermedia breviter ovato- 
oblonga, 1 mm longa, 0*8 mm lata, distante serrata, apice acuto, vix acuminate Strobili in 
apicibus ramulorum lateralium singuli dispositi, tetragoni, c. 3*5 mm longi, 1/5 mm lati, non 
ramosi; sporophylla late ovata, leviter carinata, irregulariter serrulata, apice acuto; 
megasporae c. 300 //m, pallide brunneae (humectae), scabrae rugoso-reticulataeve; micro- 
sporaec. 30 //m, verruccatae, verrucis minute spinulosis. 

Typus: Venezuela, Bolivar: Cana Mojado, Chimanta Massif, Toronotepui, 1895-1910 m, 
Steyermark & Wurdack 1079 (BM, holotype; US, isotype). 

Other specimens seen: 

VENEZUELA. Bolivar: vicinity of upper falls, Rio Tirica, Chimanta Massif, 1940 — 1950 m, 

Steyermark & Wurdack 536 (BM, US). 

Geographical range: Confined to the Chimanta Massif, Venezuela. 

67. Selaginella flexuosa Spring in Flora, Jena 21 : 197 (1838), in Martius, Fl. Bras. 1 
(2) : 122 (1840), in Bull. Acad. r. Belg. 10 : 144 (1843). Syntypes from Brazil: without exact 
locality, Swainson s.n. (K); Corcovado, Raddi s.n. (BR?); around Rio de Janeiro, 
Langsdorffs.n. (BR?); Serra de Cubatao, Guillemin s.n. (?G). The latter three not seen. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 




'VIM- SIM ( IMI \ 



Fig. 10 Selaginella scintillata Alston: Type specimen, Steyermark & Wurdack 1079 (BM). 



284 



A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 




Fig. 11 Selaginella scintillata Alston: A. Close-up of lateral (/) and median (m) leaves, x 33. B. 
Close-up of lateral (/) and axillary (a) leaves, x 24. C. Median leaves (m) and axil of lateral leaf (I) 
showing ligule, x 108. All from Steyermark & Wurdak 1079. 



Lycopodium brasiliense Desv. in Mem. Soc. linn. Paris 6 : 190 (1827), non Raddi (1825). 

Type from Brazil. 
Selaginella brasiliensis (Desv.) Spring in Flora, Jena 21 : 213 (1838), non (Raddi) A. Braun 

(1865). Type as above. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 285 

Selaginella bella Fee, Crypt. Bras. 2:100 (1873); Hieron. in Hedwigia 43:4 (1903). 

Syntypes from Brazil. 
Selaginella longicuspis Baker in J. Bot., Lond. 21 : 241 (1883). Type from Brazil. 
Selaginella macrorhyza Silveira in Bolm Comm. geogr. geol. Minas Geraes 5 : 121 (1898); 

Hieron, in Hedwigia 43 : 37 (1904). Type from Brazil. 
Selaginella brevipes sensu Baker, Fern Allies: 45 (1887), non Fee (1869) nee A. Braun 

(1867). 

Specimens seen: 

BRAZIL. Without exact locality: type as above; Glaziou 4491 (C, syntype of S. bella Fee), 4494 (C, 
syntype of S. bella Fee), 4504 (C, syntype of S. bella Fee, P), 4483 (C); Bowie & Cunningham s.n. (BM); 
Capanema s.n. (BM); Dusul 3720 (BM); Riedel s.n. (BM). Espirito Santo: Alto Limoeiro — Santa Maria, 
Brade, Pereira & Duartz 18276 (BM); Cachoeiro do Itapemirim, vargem Alta: Morro de Sul, Brade 
19359 (BM); Castelo, Braco do sul, 800 m, Brade 19211 (BM), 19214 (BM), 19310 (BM). Minas 
Gerais: Conselheiro Matta, Brade 13963 (BM); Serra do Caraca, Hoehne s.n. (BM); near Rio Preto, 
Silveira 161 (BM); Ouro Preto, de Godoy 8420 (SP); Damazio 733 (P); S. Sebastian, Damazio 732 (P); 
Silveira 170 (P); Avraial de Bento Rodrigues, Damazio 1876 (P); Bello Horizonte, Serra da Mutuca, 
Markgraf, Barreto & Brade 3573 (BM); Badini 318 (BM); Sta. Barbara do Matto Dentro, Hoehne 
5004 A (BM); S. Sebastias de Paraiso, Cachoeira do Bau, Brade & Pereira 17954 (BM). Parana: Rio 
Branco, Dusen s.n. (BM); Palmeira, Freitas 57 (BM); Villa Velha, Dusen 14856 (C), 14906 (C); 
Ypiranga, Dusen 3720 (P). Rio de Janeiro: Macgillivrav s.n. (BM), Glaziou 1645 (BM, C); Tijuca, 
Friere s.n. (BM); Alston & Brade 9012 (BM); Glaziou 7353 (BM, C, K, type of S. longicuspis); Freire 4 
(BM); serra do Cariacica, Brade 13983 (BM); Smith 1273 (BM, C); Brade s.n. (BM); Occhioni 3 (BM); 
Petropolis — Casangola Morro do Muiz, Goes & Constantino 1032 (BM); Corcovado, Peckolt & Freire 
481 (BM); Av. Niemeyer, Pilger & Brade s.n. (BM); Itatiaia, Brade 10059 (BM), 10236 (BM); 800 m, 
Brade 17221 (BM); 14529 (BM); Estrada Maromba, Vaughan Bandeira s.n. (BM); Morro Queimado, 
Brade 15722 (BM). Santa Catarina: Joinville, Mueller 133 (NY); Mueller s.n. (Rosenstock exsicc. 336; 
BM). Sao Paulo: Alto da Serra, Usteri 28a (BM, P); Luederwaldt 1658 (BM); Gehrt s.n. (BM); Santos, 
Mosen 3813 (C); Caroa da Muerte, Santos, Schreiter 1 2 (B). 

Geographical range: Endemic to Brazil. 

Notes: S. longicuspis Baker appears to differ only in size. 

68. Selaginella decomposita Spring in Martius, Fl. Bras. 1 (2): 123 (1841), Bull. Acad. r. 

Belg. 10:226 (1843). Syntypes from Brazil, Sebastianopolis, Langsdorff s.n. (?B), 

Guillermin s.n. (?BR). 
Selaginella subsegregata Baker in J. Bot, Lond. 21 : 334 (1883), Fern Allies : 59 (1887). 

Type from Brazil. 
Selaginella assurgens Baker in J. Bot., Lond. 22 : 277 (1884), Fern Allies : 78 (1887). Type 

from Brazil. 
Selaginella fusca Silveira in Bolm Comm. geogr. geol. Minas Geraes 5 : 123 (1898). Type 

from Brazil. 

Specimens seen: 

BRAZIL. Without exact locality: Glaziou 7278 (C, P), 1 1722 (C, P), 4501 (C, type of S subsegregata 
Baker), 7968 (C, syntype of S. assurgens Baker); Burchell 2238 (K, syntype of S. assurgens Baker). 
Ceara: Baturite, S. Joao, Leitz 1360 (BM). Espirito Santo: Alto Limasiro, Brade, Altaniro & Apparici 
18129 (BM); Castelo, Forno Grande 1000 m, Brade 19291 (BM): Cachoeiro do Iterpemirim, Vargem 
Alta, Morro de Sal, Brade 19360 (BM). Minas Gerais: Serra do Rio Preto, Silveira 162 (BM). Parana: 
Ypiranga, Dusen 8322 (BM); Lucena, Wielewski 1 1 (NY). Rio Grande do Sul: Serra do Estrella, 700 m, 
Brade 1 8657 (BM). Rio de Janeiro: Glaziou 52 1 5 (C, K, syntype of S assurgens Baker), 450 1 (K), 5637 
(C, K, P, syntype of S assurgens Baker), 7768 (K): Itatiaia, Brade 10190 (BM), 10238 (BM); 
Maromba, Brade 14532 (BM); Petropolis l Independencia\ Alston & Lutz 273 (BM): Cachamba Road, 
Petropolis, Glaziou s.n. (BM, P); St Antonia de Finbe, Brade & Luis 1 1663 (BM): Serra dos Orgaos, 
Gardner 76 (K, syntype of S. assurgens): Guillemin 962 (P); Terezopolis, Brade s.n. (BM): Terezopolis, 
Cachoeira Teroz, Lutz 2087 (BM). Santa Catarina: Laranjeira mts. Vie 129 (P): Queimadas, Spannagel 
196 (NY). Sao Paulo: Usteri s.n. (BM); Alto da Serra, Luederwalt 1656 (BM): Gehrt 4396 (BM): Usteri 



286 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

25a (P), 21038 (P); Lorena, Luederwaldt s.n. (BM); Serra do Paranapiacaba, Brade 15716 (BM); 
Apiahy,ScAw0cte2385(P). 

Geographical range: Endemic to Brazil. 

Notes: S. subsegregata Baker and S. assurgens Baker are luxuriant specimens, S. fusca 
Silveira has rather more acute leaves than average S. decomposita but is still within the 
range of the species. It is closely allied to the West Indian S. substipitata Spring. 

69. Selaginella duidae A. C. Smith in Bull. Torrey bot. Club. 58 : 312 (1931). Type from 
Venezuela, Bolivar: Laterite Valley, Savanna Hills, Duida, 1320 m, Tate 839 (NY, 
holotype; BM, isotype). The only specimen seen. 

Geographical range: Confined to Mt Duida. 

70. Selaginella euclimax Alston ex Crabbe & Jermy in Fern Gaz. 11 : 259 (1976). Type from 
Colombia, Cauca: W of Tambo, 2300 m, Haught 5220 (BM, holotype; COL, US, 
isotypes). 

Specimens seen: 

COLOMBIA. Antioquia: Rio Guapope, 1 5-1 800 m, Kalbreyer 1457 (B, BM). Cauca: type as above. 

Geographical range: Confined to Colombia. 

Notes: This species resembles S. tarapotensis (No. 99), which is prostrate rather than erect, 
and has a longer arista on the median leaves. S. pearcei (No. 34) is also very similar, but it has 
broader lateral leaves with acute apices, very short aristae, and is usually less branched 
below. On the herbarium label in BM of Kalbreyer 1457 the altitude is given as 
5-6000 metres, which cannot be the case; the error was perpetuated in Crabbe & Jermy 
(1976). 

71. Selaginella vernicosa Baker in Timehri 5 : 220 (1886), in Trans. Linn. Soc. Lond. (Bot.) 
52 : 295 (1887), Fern Allies : 78 (1887). Type from Venezuela, Bolivar: SE slopes of Mt 
Roraima, 4 Our House', 1622 m, im Thurn 266 (K, US). 

Selaginella vernicosa var. oligoclada Baker in Trans. Linn. Soc. Lond. (Bot.) 2 : 295 (1887), 

Fern Allies : 78 (1887). Type from Venezuela. 
Selaginella vernicosa var. simplicifrons Jenman, Ferns & Fern Allies W. Indies : 401 (1908), 

in synonymy. 

Specimens seen: 

VENEZUELA. Bolivar: type, as above; Mt Roraima, summit, 2700—2740 m, McConnell&Quelch 574 
(BM, NY); Tate 416 (BM, NY); Steyermark 57796 (US), 58016 bis & ter (US), 58889 (BM); SE slopes 
'Our House' 1622 m, im Thurm 381 (BM, type of S. vernicosa var. oligoclada Baker); above Rondon 
Camp, 2070 m, Tate 466 p.p. (NY); Chimanta Massif, Central section, 1940 m, Steyermark & 
Wurdack 404 (BM). 

Geographical range: Confined to La Gran Sabana. 

72. Selaginella vestiens Baker in J. Bot., Lond. 21 : 97 (1883), Fern Allies : 43 (1887). Type 
from Brazil, Goias: Morro de Canto Gallo, Burchell 7006 (K). 

Selaginella cladostachya Baker in J. Bot., Lond. 21 : 97 (1 883). Type as above. 
Selaginella erythrospora Silveira in Com. Geogr. Geo). Est. Minas Geraes 5 : 126 (1898). 
Type from Brazil. 

Specimens seen: 

BRAZIL. Bahia: Serra Sincora, 1400 m, Vie 7298 (BM). Goias: type, as above. Minas Gerais: St 
Hilaire 913 (P); Serro da Cipo, Silveira 156 (BM, type of S. erythrospora A. Silveira); Strada Pila, 
Barret 58 1 & Brade 144404 (BM); near Aguas Santas, Serra de S. Jose, Silveira 2622 (P); Campos de S. 
Sebasiao, Damazio 1882 (BM, P): Christias, near Corriga dois Puntes, Diamantina, Mexia 5832 (BM); 



SELAGINELLA IN TROPICAL SOUTH AMERICA 287 

Serra do Rio Grande, 1260 m, Diamantina, Mexia 5799a (BM); Consulheiro Matta, Brade 13962 
(BM). Rio de Janeiro: Morro de Cubicado, Petropolis, Glaziou 1 1 723 (BM, C, P). 

Geographical range: Confined to south-east Brazil. 

Notes: Selaginella cladostachya was described by Baker at the same time as S. vestiens and 
from the same gathering made by Burchell. The variation in leaf length, width and shape of 
apex, and degree of density of cilia varies within the range of the species, and in our opinion 
these two entities cannot be distinguished. 

73. Selaginella anisoclada Alston ex Crabbe & Jermy in Fern Gaz. 11 : 255 (1976). Type 
from Venezuela, Aragua: above Guamitas Parcque Nacional, 900 m, Alston 58 14 (BM). 

Specimens seen: 

VENEZUELA. Aragua: type as above. 

COLOMBIA. Magdalena: Sierra del Libano, Santa Marta, 1650 m, Smith 2242 (BM, NY, US); Santa 

Marta, Smith 2567 p.p. (BM). 

Geographical range: Confined to the coastal ranges of Venezuela and Colombia. 

Notes: The species is distinct in its distant arrangement of both median and lateral leaves, 
which are almost isomorphic on the ultimate branches. It varies in stature of plant and 
degree of branching, a character obviously affected by the moistness of, and competition in, 
the habitat. It is most similar to S. porphyrospora (No. 44), but there the lateral leaves are 
larger, lanceolate and acute. S. lychnuchus (No. 47) can be similar in habit, but that species 
has dimorphic sporophylls and large, oblong, lateral leaves, and long, aristate, median 
leaves. 

74. Selaginella microphylla (Kunth) Spring in Bull. Acad. r. Belg. 10 : 234 (1843), in Mem. 
Acad. r. Sci. Lett. Belg. 24:88 (1850); Baker, Fern Allies :41 (1887), p.p. excl. pi. 
guatemalensis. Type from Colombia, Cauca: Qulcace, Bonpland s.n. (BM). 

Lycopodium microphyllum Kunth, Nov. Gen. Spec. 1 : 37 (1816), Syn. PI. Itin. Orb. Nov. 

Humb.Bonpl. : 96 (1822); Spreng, Syst. Veg.4: 15 (1827). Type as above. 
Selaginella thujaefolia Spring in Martius, Fl. Bras. 1, pt 2 : 120 (1840), in Bull. Acad. r. Belg, 

10 : 140 (1843). Type from Uruguay. 
Selaginella jamesoni Baker in /. Bot., Lond. 21 : 97 (1883), Fern Allies : 42 (1887). Syntypes 

from Ecuador. 
Selaginella microphylla f. communis Sodiro, Rec. Crypt. Vase. Quito : 92 (1883), nom. nud. 
Selaginella microphylla f. culcitrella Sodiro, Rec. Crypt. Vase. Quito : 92 (1883). Type from 

Ecuador. 
Selaginella schmidtchenii Hieron. in Hedwigia 43 : 40 (1904). Type from Colombia. 

Specimens seen: 

VENEZUELA. Merida: Moritz 369 (BM); Merida, 1 800 m, Alston 6703 (BM). 

COLOMBIA. Without precise locality: Schmidtchen s.n. (B, type of 5". schmidtchenii Hieron.; BM). 
Antioquia: near Antioquia, 550 m, Barkley, Klevens & Gutierrez 1 7c377 (BM, COL). Cauca: Qulcace, 
104 hexap., Bonpland s.n. (BM, type collection of Lvcopodium microphyllum). Cundinamarca: 
Choachi, 1800 m, Lindig 1519 (BM); Camino de Villavicencio, Dawe 273 (US). Huila: 3 km SE of 
Neiva, 510 m, Little 9475 (COL); 6 km N of Neiva, 450 m, Little 8945 (COL). Narino: Rio Guaitara, 
1500 m, Lehmann 111 (NY); Pasto, 2500 m, Espinosa 2750 (BM). Tolima: Chicoral, 450 m, Haught 
6386 (BM, COL); Gualandaya, between Espinal and Ibague, 700 m, Alston 7704 (BM). 
ECUADOR. Without precise locality: Spruce 4787 (BM, CGE, NY). Azuay: near Cuenca, Rose 22869 
(NY, US). Chimborazo: Canon of Rio Chanchan near Huigra, 1200-1350 m Camp E 3205 (BM); near 
Huigra, Rose 23959 (NY); Chambo, 2800 m, Asplund 5952 (S). Pichincha: Jameson 280 (BM); 
Machbnagara, Sodiro 30 (NY); near Quito, Couthouv 84 (NY), 2850 m, Mille s.n. (NY); below 
Guapulo, near Quito, 2500 m, Asplund 6708 (S); Chaupi-sagcha, Pululagua, 1800 m Bell 444 (BM); 
Panecillo, Quito, 2900 m, Asplund 6071 (S). Santiago-Zamora: Tambo Chontal to Tambo Consuelo, 
1710-2400 m Camp E 1586 (BM). Tungurahua: Banos, 1400 m, Asplund 19904 (S). 



288 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

PERU. Without precise locality: Mathews s.n. (BM, NY). Cuzco: San Miguel, Urubamba valley, 
1 800 m, Cook & Gilbert 1 782 (US); Quellounu, 950 m, Buez s.n. (B). 

BOLIVIA. Without precise locality: RusbylWl (NY). Cochabamba : Cochabamba, Bang 1097 (BM, 
NY); Sailapata, Ayopaya, 2700 m, Cardenas 3234 (BM); on rock, lomo de San Pedro, Botanic Garden 
2520 m, Hermann 24600 (BM). La Paz: Sorata, Larecaja, 2250 m, Williams 1404 (NY); 2400 m, 
Rusby 456 (NY); Yungas, Jay (NY), a[bis]d (NY); Sorata (E of L Titicaca), 2400 m, Brooke 6649 
(BM). Santa Cruz: Cortes, 1800 m, Williams 1403 (NY); Cerro La Negra, 1800 m, Steinbach 8186 
(NY); Cerro Tres Cruzas, 1 500 m, Steinbach 8165 (NY). 

BRAZIL. Without exactly locality: Glaziou 7967 (C). Minas Gerais: Caldas, Mosen 2025 (BM, NY, P). 
Rio Grande do Sul : Antonio Prado, Dutra 1 16 (BM); Rio Pardo, Fazenda Leitao, Jurgens L7 (BM, 
NY); Bohmerbach, Jurgens L28 (BM, NY). Sao Paulo: Serra de Botucatu, Hoehne & Gehrt 33529 
(BM); campos do Jordao, Porto 3113 (BM); Pedra do Bau, Sao Bento do sa pucai, 1900 m, Leitz 3929 
(BM). Santa Catarina: Serra do Oratorio, Vie 297 (BM, P); Braco serafim, Itajai, 200 m, Reitz 3016 
(BM). 

Geographical range: From Venezuela along the Andes to Argentina, and east to southern 
Brazil, Paraguay and Uruguay. 

Notes: Selaginella schmidtchenii was described from an abnormal specimen with no further 
data than 4 Nova Granada, 1882'; in our opinion it is a form of S. microphylla; the curling up 
of the lateral leaves might be caused by drought or frost. Similar specimens have been 
collected elsewhere in Colombia by Haught 6386 (BM) and Alston 7704 (BM). 

75. Selaginella schultesii Alston ex Crabbe & Jermy in Am. Fern J. 63 : 141 (1973). Type 
from Colombia, Amazonas: Rio Popayaca, tributary of R. Apaporis, Schultes & Cabrera 
15561 (US, holotype; BM, isotype). 

Specimens seen: 

COLOMBIA. Amazonas: type as above; Raudal Yayacopi, La Playa, 240 m, Schultes & Cabrera 

15417 (US), 16205 (US), 16898 (BM, US). Vaupes: Cachoiera Miriti, Rio Kanari, 250 m, Schultes & 

Cabrera 14411 (COL, US); Cerro Isibukuri, Rio Kananari, 250-700 m, Schultes & Cabrera 13314 

(COL, US); Rio Apaporis, Cachivera de Jirijirimo y alrededores, 250 m, Schultes & Cabrera 12456 

(COL, US); Rio Apaporis, Raudal de Jirijirimo 270 m, Schultes & Cabrera 14554 (COL), 14598 

(COL). 

Geographical range: Confined to Colombia. 

Notes: Schultes notes (in sched.) that this plant is known to the Makuna Indians as 4 wee-wee- 
a-ree' suggesting that it may have had a local use. S. homaliae (No. 95) is similar in habit, but 
has more oblong lateral leaves, usually twice as long as broad, which are rounded at the apex 
and almost without cilia. S. applanata (No. 98) is again similar, but has more elongated and 
even larger leaves (up to 2*5 mm). 

76. Selaginella microdonta A. C. Smith in Bull. Torrey hot. Club 58 : 313 (1931). Type from 
Venezuela, Amazonas: on slope of Ridge 24 and summit of Mt Duida, 1800 m, Tate 509 
(NY, holotype; BM, isotype). 

Specimens seen: 

VENEZUELA. Amazonas: types as above. Bolivar: Ptari-teoui, 2885-2405 m, Steyermark 59563 

(BM,US). 

Geographical range: Mountains of southern Venezuela. 

Notes: This species resembles S. rotundifolia Spring of the West Indies, but the median leaves 
are narrower and more acuminate. S. ovifolia subsp. philipsonii (No. 83) differs in having 
distinctly ciliate lateral leaves. 

77. Selaginella jungermannioides (Gaudich.) Spring in Bull. Acad. r. Belg. 10 : 143 (1843); 
Baker, Fern Allies : 53 (1 877) Type from Brazil, Rio de Janeiro, Gaudichaud s.n. (K). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 289 

Lycop odium jungermannioides Gaudich. in Freyc, Voyage Autour Monde, Bot. 286 ['826'] 

( 1 828). Type as above. 
Lycopodium marginatum sensu Raddi, PL Bras. Nov. Gen. : 82 (1825), excl. syn. 

Specimens seen: 

BRAZIL. Without exact locality: Glaziou 4675 (C), 7493 (C), 7965 (C, P), 12295 (C). Espirito Santo: 
Rio Mutuni, Luetzelburg 9942 (M). Rio de Janeiro: type as above; Raddi s.n. (BM); Mosen s.n. (M); 
Tijucca, Usteri s.n. (BM); Alston & Brade 9004 (BM); Brade 1 003 1 (BM); Corcovado, Rudio s.n. (BM); 
Luetzelburg 9960 (M); Serra Estrella, Luetzelburg 7386 (M), 7875 (M); Serra dos Orgaos, Luetzelburg 
7045 (M); Una Grande, Rose & Russell 20349 (NY); St Antonio de Finba, Brade & Luis 11672 (BM). 

Geographical range: Confined to eastern Brazil. 

78. Selaginella arenaria Baker in J. Bot., Lond. 21 : 82 (1883), Fern Allies: 40 (1887); 
Alston in Reprium Spec. nov. Regni veg. 40 : 317 (1936), p.p. quoad spec, typic. Type 
from Brazil, Amazonas: On Rio Vaupes at Panure, Spruce 286 1 (K). The only collection. 

Geographical range: West Brazil. 

Notes: There is a note on the type sheet at K, in what appears to be Spruce's handwriting, 
stating: 4 I believe some barren specimens of this were mixed up with No. 2535; the two are 
however quite distinct.' Specimens under this collecting number have been distributed to 
several herbaria; but so far the type is unique and the other specimens do not match any 
hitherto described species. They are described as S. calceolata (No. 1 18, p. 3 14). 

79. Selaginella brevifolia Baker in J. Bot., Lond. 21 : 83 (1883), Fern Allies :41 (1887). 
Alston in Reprium Spec. nov. Regni veg. 40 : 312 (1936). Type from Brazil: Spruce 2547 
(BM, CGE, K). 

Selaginella cryptogaea Baker in J. Bot., Lond. 21 : 98 (1883), Fern Allies : 44 (1887). Type 
from Brazil, Amazonas: on rocks by the Janarate Cachoeira, near Panure, Spruce 2547 (K 
holotype; BM, CGE, isotypes). 

Specimens seen: 

VENEZUELA. Amazonas: Rios Pacimoni-Yatua, Casiquiare, 1 10-130 m, Maguire, Wurdack & 
Bunting 37468 (BM). Bolivar: Saltode Para, Medio Caura, 300 m, Williams 1 1385 (BM). 
COLOMBIA. Huila: 1 km east of Neiva, 510 m, Little 9250 (COL). Vaupes: Rio Vaupes, 75 m, 
Schultes & Cabrera 13866 (BM); Cerro de Tipiaca, between Mitu and Javarete, Schultes & Cabrera 
19319 (US); Cerro Yapoboda, Rio Kuduyari, 450 m, Schultes & Cabrera 14182 (COL, US), 14215 
(COL, US). 

PERU. San Martin: Boqueron between Tingo Maria and Pucallpa400 m, A I 'lard 22124 (BM). 
BRAZIL. Amazonas: type, as above; near San Gabriel da Cachoeira, Rio Negro, Spruce 2328 (BM, 
CGE); Schultes & Lopez 9171 (BM); Piedra de Cocui, Rio Negro, Schultes & Lopez 9428 (BM); San 
Felipe, Luetzelburg 22129 (M); Serra Wabeesee, below Bela Vista, Rio Uaupes, Schultes & Pires 9147 
(BM); near Panure, Rio Uaupes, Spruce 2905 (BM, K, type of S. cryptogaea Baker). 

Geographcal range: Venezuela, Colombia, Peru and the western part of Brazilian 
Amazonas. 

Notes: This species is very closely allied to S. cordifolia (Desv.) Spring from Cuba, 
Hispaniola and Porto Rico, but that species has longer, aristate, median leaves and is often 
flagelliferous. S. muscosa (No. 88) may be confused with some forms of S. brevifolia, but the 
lateral leaves are denticulate and the median leaves have longer aristae. Maguire, Wurdack 
& Bunting 37468 has distinct almost triangular leaves on the main stem. Allard 22124 is a 
sterile specimen with median leaves that have oblique bases with an auricle on the outer side; 
it is close to this species but may be found not to be conspecific when more 
material has been studied. 

80. Selaginella sandwithii Alston, sp. nov. (Figs 1 2 & 13) 

Species heterophylla ex affinitate S. deltoidei sed habitu erecto et rhizophoris ad basin 
limitatis differt. 



290 



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Fig. 12 Selaginella sandwithii Alston: Type specimen, Sandwith 1 277 (BM). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



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Fig. 13 Selaginella sandwithii Alston: A. Close-up of lateral (/) and axillary {a) leaves, x 15. B. 
Close-up of median (m) and lateral leaves; note laminal teeth (unicellular papillae) on upper 
surface of lateral leaves; x22. C. Median leaf, lower surface, x460. D. Lateral leaf, upper 
epidermis with unicellular papillae, x 143. All from Sandwith Mil. 



Planta e basi repente ascendens vel erecta; caulibus glabris, 6-14 cm altis, basi 1 mm in 
diametro, e basi ramosis vel parte simplici usque ad 10 cm longa; rhizophoris ad tertiam 
partem basin versus restrictis; parte ramosa ambitu plerumque irregulariter ovata, ramis c. 
1 cm inter se distantibus, pinnatis ambitu auguste ellipticis, usque ad 2*5 cm longis. Folia 
heteromorpha vel in parte simplici basin versus subaequalia; folia lateralia oblique patentia, 
crebre disposita vel leviter imbricata, ovato-deltoidea 2 mm longa, 125 mm lata, acuta; in 
epidermide superiore marginem versus papillis unicellularis praedita; semifacie superiore 
semi-ovatodeltoidea, basi rotundato-cordata breviter serrulato-ciliolata distanter minutis- 
simeque serrulata, aliter integra; semifacie inferiore semi-oblongo-deltoidea, basi truncata, 
apice cuneato et distanter minutissimeque serrulato, alibi integra; folia axillaria ovato- 



292 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

deltoidea, quam folia lateralia paullo minoria; folia intermedia ovata, c. 1 mm longa, 0*7 mm 
lata, serrata, valde albo-marginata basi leviter inaequilateralia, apice acuminato-aristato; 
arista dimidio breviore lamina. Strobili ad apicibus ramulorum omnium singulariter 
dispositi, subtetragoni, 3 mm longi, 1*5 mm lati, non ramosi; sporophylla anguste ovata, 
carinata, albomarginata, serrata, acuminata; megasporae non visae; microsporae c. 37 jum 
luteae (humectae) verrucosae verrucis minute spinulosis. 

Typus: Guyana: Essequibo, Potaro River, ascent from Tukeit to Kaieteur savannah, 
150-270 m, Sandwith Mil (BM, holotype; K, isotype). 

Other specimens seen: 

SURINAM. Saramacca: Tafelberg, Maguire 24336 (BM, NY), 24553 (BM, NY), 24772 (BM, NY). 
GUYANA. Essequibo: Kaieteur gorge, Tutin 536 (BM); Sandwith 1501 (BM); Kaieteur Plateau, 
Maguire & Fanshawe 23416 (BM). 

Geographical range: Sandstone area of Surinam and Guyana. 

Notes: Alston was long puzzled by this species and referred some specimens to S. flagellata 
(No. 41) and others to S. muscosa (No. 88). The former differs by its flagelliform apices and 
microspores with elongate papillae; the latter by its prostrate habit. S. brevifolia (No. 79) is 
also similar, but differs by its ciliate lateral leaves. We are retaining this as a new taxon with 
the belief that, as more material becomes available, it may be merged with S. deltoides A. 
Braun, a segregate of S. muscosa (see notes on p. 298). 

81. Selaginella valdepilosa Baker in /. Bot., Lond. 21 : 82 (1883), Fern Allies : 40 (1887); 
Knox in Trans, bot. Soc. Edinb. 35 : 250 (1950). Type from Guyana: Essequibo: ravines 
near the Kaieteur Savannah, Jenman 1484 (K, holotype; BM, NY, isotypes). 

Two subspecies are distinguished: 

subsp. valdepilosa. 

Specimens seen: 

GUYANA. Essequibo: type as above; on precipices of Kaieteur Gorge. 360 m, Sandwith 1435 (BM, K); 

Waratula, Potaro R., Tutin 480 (BM); Maguire & Fanshawe 23415 (BM). 

Geographical range: Confined to Guyana. 

Notes: Two specimens in BM (Brazil: Rio Grande do Sul: Montenegro, S. Salvador, 
Sehnem s.n. [Hb Dutra 1233]) are very near this species morphologically, but need further 
investigation. 

subsp. tricholoma Jermy & Rankin, subsp. nov. (Fig. 14) 

Planta subsp. valdepilosae similis sed foliis angustioribus, plerumque inter se distantibus 

differt. 

Planta omnino repens prostrataque; caulibus glabris, flexuosis, subfiliformibus, in sicci- 
tate pallide stramineis, c. 12 cm longis, distanter alternatim pinnatis nonnunquam bipin- 
natis, rhizophora ubique gerentibus; ramis c. 10 mm inter se distantibus, usque ad 25 mm 
longis, furcatis. Folia ubique heteromorpha; folia lateralia patentia, paullum distantia vel 
subcontigua, elliptico-oblonga, 1 mm longa, - 75 mm lata, obtusa, fere aequilateralia, 
utrinque ciliata; semifacie superiore late semi-elliptico-oblonga; semifacie inferiore semi- 
el liptico-oblonga; folia axillaria aequilateralia; folia intermedia elliptico-orbiculata, basi 
rotundata, 07 mm longa, 060 mm lata, apice obtusa, ubique ciliata, oblique patentia; 
strobili non visi. 

Typus: Guyana: Essequibo: vertice montis Roraima, 2700-2740 m, Steyermark 58016 
(BM, holotype; US isotype). 

Geographical range: Known only from the type collections from Mt Roraima. 

Notes: This species also resembles S. rhodostachya (No. 86), which has more ovate lateral 
leaves. Material under the type number, labelled 58016 bis (BM, US) is a mixture of this 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



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Fig. 14 Selaginella valdepilosa subsp. tricholoma Jermy & Rankin: A. Type specimen, 
Steyermark 58016 (BM). B. Close-up of lateral leaf, x 40. C. Close-up of axillary leaf, x 25. D. 
Close-up of median leaves, x 40. 



294 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

subspecies and S. vernicosa (No. 71), and is best treated as a paratype. Alston was inclined to 
treat this taxon as a species; we feel it is very close to S. valdepilosa s.str. and at the moment 
prefer to give it subspecific rank, but as more material is collected it may even prove to be a 
growth form of that species. 

82. Selaginella tragi I lima Silveira in Bolm Comm. geogr. geol. Minas Geraes 5 : 127 
(1898); Hieron. in Hedwigia 43 : 12 (1903). Type from Brazil: Serra de S. Jose d'El-Rei, 
Silveira 2622 (Herb. Com. Geog. e Geolog. Civitatis Minas Geraes). 

Specimens seen: 

BRAZIL. Minas Gerais: type, as above; S. Sebasitas de Paraiso, Pereira, Brade & Barbosa 17953 (BM); 

Serra do Lenheiro, Silveira 149 (BM). 

Geographical range: Endemic to Brazil. 

83. Selaginella ovifolia Baker in J. BoL, Lond. 2 : 90 (1884), Fern Allies : 68 (1887); Alston 
in Bull. Br. Mus. nat. Hist. (Bot.) 1 : 45 (1952) & 244 (1955). Type from Puerto Rico, 
Schwanecke s.n. (K). 

Subsp. philipsonii Jermy & Rankin, subsp. nov. (Figs 1 5 & 1 6) 

Subspecies a subsp. ovifolia foliis lateralibus minus acutis, longe ciliatis, foliis intermediis 

majoribus oblongioribusque differt. 

Planta omino repens et prostrata; caulibus glabris, filiformibus, pallide luteo-viridis, 
12 cm longis, distanter alternatim pinnatis, nonnunquam bipinnatis; rhizophora ubique 
gerentibus; ramis c. 1 cm inter se distantibus, usque ad 2 cm longis. Folia ubique hetero- 
morpha; folia later alia patentia, distantia, ovato-rotundata, 1 mm longa 0'75 mm lata, 
obtusa, fere aequilateralia; semifacie superiore, longe ciliata fere ad apicem; semifacie 
inferiore, basi integra, et apicem versus serrata; folia axillaria similia lateralibus; folia 
intermedia oblongo-elliptica, distantia, latere exteriore integra, basi rotundata et latere 
interiore serrata, basi cuneata, 0*6 mm longa, 0*45 mm lata, apice subacuto. Strobili non visi. 

Typus: Colombia, Meta/Vaupes: Rio Guapaya, Macarena, 450 m, Philipson, Idrobo & 
Fernandez 1607a(BM, holotype). 

Other specimens seen: 

COLOMBIA. Meta: Margen del Rio Guayabero, 10 km abajo de Cano Lozada, 350 m, Pinto & 
Bischler 239 (COL); Sierra La Macarena, 'Cano Veinte', 500 m, Idrobo 4837 (BM). Norte de 
Santander: Catatumbo, Campo Oru y alrededores, 350-500 m, Bischler 2397 (COL); Catatumbo, 
Campo Tibu, 200 m, Bischler 2493 (COL). 

Geographical range: confined to the Macarena Mts, Colombia. Subsp. ovifolia occurs in 
Central America and the West Indies. 

Note: Alston treated this as a new species, but its similarity and presumed affinity to S. 
ovifolia Baker is so great that we have designated it as a subspecies of that species. It has 
lateral leaves which are obtuse with occasionally a mucronate apex, and median leaves 
which are less ciliate and more oblong than in the type subspecies. The specimens are sterile 
but we have confidence in designating it as a new taxon. 

84. Selaginella cabrerensis Hieron. in Hedwigia 43:29 (1904). Type from Colombia, 
Tolima: Rio Cabrerea, near Colombia, 500-1000 m, Lehmann 6406 (K, US). 

Selaginella daguensis Hieron. in Hedwigia 43 : 49 (1904). Type from Colombia. 

Specimens seen: 

COLOMBIA. Cundinamarca: Pandi, 900 m, Andre 1817 (BM, NY). Meta: Serra de la Macarena, 
900 m, Philipson 2388 (BM). Tolima: type as above: Chichoral, 450 m, Haught 6333 (BM), 6439 (BM, 
COL). Valle de Cauca: Rio Dagua near Buenaventura, 0-300 m, Lehmann 4462 (K, type collection of 
S. daguensis Hieron.). 

Geographical range: Confined to Colombia. 






SELAGINELLA IN TROPICAL SOUTH AMERICA 



295 






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Fig. 15 Selaginella ovifolia subsp. philipsonii Jermy & Rankin: Type specimen, Philipson, 

Idrobo and Fernandez 1 607a (BM). 



296 



A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 




Fig. 16 Selaginella ovifolia subsp. philipsonii Jermy & Rankin: A. Close-up of lateral leaves, 
x 24. B. Close-up of median leaves, x 88. All from Philipson, Idrobo & Fernandez 1607a. 



Notes: Hieronymous relates S. cabrerensis to S. microphylla, and S. daguensis to S. apus, 
and does not compare them with one another. We can see no difference except that S. 
daguensis is more erect. 

85. Selaginella tuberculata Spruce ex Baker in J. Bot. Lond. 21 : 83 (1883), Fern Allies : 40 
(1887); Alston in Reprium Spec. nov. Regni veg. 45 : 3 12 (1936). Type from Brazil: Panure 
Falls, Rio Uapes, Spruce 2940 (K, holotype; BM, CGE, isotypes). 

Specimens seen: 

SURINAM. Saramacca : Tafelberg, Geyskes 998a (BM); Maguire 24339 A (BM). 

GUYANA. Essequibo: Kaieteur, Jenman s.n. (BM, NY), Waratuk, Tutin 480 (BM); between Tukeit 

and the Kaiteur savanna, Sandwith 1278 (BM, K): Guppy 168 (BM). 

BRAZIL. Amazonas: Type as above. 

Geographical range: Sandstone area of Surinam and Guyana to northern Brazil. 

Notes: This species differs from S. rhodostachya (No. 86), with which Jenman confused it, by 
its ciliate lateral leaves with tubercles on the lower half of the upper surface. 

86. Selaginella rhodostachya Baker in Timehri 5 : 221 (1886), in Trans. Linn. Soc. Lond. 
(Bot.) 2:296 (1887), Fern Allies: 112 (1887). Type from Guyana, Essequibo: foot of 
Kaieteur Falls, Jenman 1 48 (K, holotype; BM, isotype). 

Selaginella brachyclada Baker, Fern Allies : 45 (1 887). Type from Guyana. 
Selaginella diminuti folia Jenman in Gdnrs' Chron. Ill, 2 : 99 (1887). Type from Guyana. 
Selaginella brevispicata Hieron. ex Ule in Bot. Jb. 52:49 (1915), nomen nudum; ex 
Bautista in Acta AmazonicaA : 19 (1974). Type from Venezuela. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 297 

Specimens seen: 

GUYANA. Essequibo: type as above; ibid, loc., Jenman 1481 (BM, K, type of 5". brachyclada Baker; 

NY, type of S. diminutifolia Jenman); path to top of Kaieteur Falls, Jenman s.n. (NY); Roraima, 

Warrington, Burr as, Woodhams & Edwards s.n. (K). 

VENEZUELA. Bolivar: Paritepui, 2130 m, Steyermark 59836 (BM); Chimanta Massif, Torono-tepui, 

Steyermark & Wurdack 1181 (BM); Mt Roraima, 2255-2620 m. Steyermark 58753 (BM, US); 

Abbensetts 1A (BM, K); Jenman s.n. (NY); Rondon Camp, 2070 m, Tate 466 p.p. (BM, NY); Vie 

8491 (BM, K; type of S. brevispicata Hieron.); im Thurn s.n. (BM); summit of Roraima, 270-2740 m, 

Steyermark 589 \ 6 {BM). 

Geographical range: Confined to southern Guyana and the Roraima massif. 

Notes: Specimens which match exactly the type have not been collected since Jenman's 
original gathering, perhaps because few botanists reach the foot of the Kaieteur. Some other 
specimens bear rhizophores throughout, and some have slightly narrower leaves. We believe 
the variation is such that there is a cline to S. brevispicata on the upper slopes of Roraima. 
Baker has dated the British Museum (Natural History) copy of his Fern Allies 'July T and 
the Gardeners' Chronicle appeared on23July 1887. 

87. Selaginella potaroensis Jenman in Gdnrs' Chron. Ill, 2 : 154 (1887), Ferns Brit. W. 
Indies : 396 (1909). Type from Guyana, Essequibo: Kaieteur Falls, Jenman 1818 (NY, 
holotype; BM, K, isotypes). 

Specimens seen: 

GUYANA. Essequibo: type as above; Mt Roraima, Waruma Trail, 1380 m, Persaud 124 (UG); 

between Tukeit and Kaieteur Falls, Guppv 164 (BM); Maguire & Fanshawe 23083 (BM); Sandwith 

1275 (BM,K). 

SURINAM. Saramacca : Arrowhead Basin; Tafelberg, Maguire 24497 (BM, NY). 

VENEZUELA. Amazonas: Cerro Duida, 1096-1520 m, Steyermark 57999 (BM). Aragua: El Cumbre, 

Parque Nacional, 1100 m, Alston 5318 (BM); La Regresiva, km 25, Parque Nacional, 1000 m, 

Williams & Alston 1 70a (BM). Lara: Barquisemento, Moritz s.n. (BM). 

Geographical range: Surinam to Venezuela. 

88. Selaginella muscosa Spring in Martius, Fl. Bras. 1 (2) : 120 (1840). Type from Brazil: 
Sebastianopolis, Luschnatt s.n. (B ?; not seen). 

Lycopodium brasiliense Raddi, Pi Bras. Nov. Gen. : 82 (1825). Type from Brazil. 
Lycopodium crassinervium Desv. in Mem. Soc. linn. Paris 6 : 190 (1827). Type from Brazil. 
Lycopodium pallidum Beyr. ex Gaudich. in Freyc, Voyage Autour Monde, Bot.: 285 (1828), 

nomen abortivum. 
Lycopodium albidulum var. majus Hook. & Grev. in Bot. Misc. 2 : 399 (1831), p.p. Types 

from Brazil. 
Selaginella crassinervia (Desv.) Spring in Martius, Fl. Bras. 1 (2): 119 (1840). Type as 

above. 
Selaginella apus var. tetragonostachya Spring in Martius, Fl. Bras. 1 (2) : 119 (1840). Type 

from Brazil. 
Selaginella polysperma Spring in Bull. Acad. r. Belg. 10 : 138 (1843). Type from Brazil. 
Selaginella beyrichii A. Braun, Index Sem. Hort. bot., Beri Appendix 1857: 16 (1857). Type 

from cultivated material. 
Selaginella anocardia A. Braun in Annls Sci. nat. (Bot.) V, 3 : 290 ( 1 865). Type from Brazil. 
Selaginella brasiliensis (Raddi) A. Braun in Annls Sci. nat. (Bot.) V, 3 : 290 (1865), non 

(Desv.) Spring (1838). 
Selaginella deltoides A. Braun in Annls Sci. nat. (Bot.) V, 3 : 287 (1865); Alston in Reprium 

Spec. nov. Regni veg. 45 : 3 12 (1936). Syntypes from Brazil. 
Selaginella brevipes Fee, Crypt. Bres : 226 (1869), non A. Braun (1867). Type from Brazil. 
Selaginella trifurcata Baker in J. Bot., Lond. 21 : 98 (1883), Fern Allies : 43 (1887); Alston in 

J. Bot., Lond. 72 : 38 (1934); Knox in Trans, bot. Soc. Edinb. 35 : 278 (1950). Type from 

Brazil. 



298 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Selaginella niederleinii Hieron. in Bot. Jb. 22 : 418 (1896). Type from Argentina. 
Selaginella feei Hieron. in Engl. & Prantl, Nat. Pflanzenf. 1 (4): 713 (1901). Type from 

Brazil. 
Selaginella brasiliensis var. crassinervia Hieron. ex Bonap., Notes Pterid. 2 : 137 (1915), 

nom. nud. Syntypes from Brazil. 
Selaginella humilis sensu Hieron. in Hedwigia 58 : 32 1 (19 1 7), non Jenman (1 897). 

Specimens seen: 

GUYANA. Berbice: Sierra Acari, Manicole bog, Guppy 336a (BM). Essequibo : Kaieteur gorge, below 
the falls, Sandwith 1478 (BM, K); Tutin 535 (BM); Kaieteur savannah, 300 m, Tutin 502 (BM). 
VENEZUELA. Delta Amacuru : Imataca Mts, Thompson 2 1 (BM). 

COLOMBIA. Caqueta : Solano, 8 km SE of Tres Esquinas, on Rio Caqueta below mouth of Rio 
Orteguaza, 200 m, Little 9583 (COL). Magdalena: Mts above Hacienda Cincin, Santa Marta, 2100 m, 
Foster & Smith 1392 (COL). Narino: between Paramo and Barbacoas, 500 m, Alston 8468 (BM). 
Vaupes: Cerro de Mitu, 250 m, Schultes & Cabrera 13931 (COL, US); Raudal Guacuraba between 
Mitu and Javaraete, Schultes & Cabrera 19268 (US); Rio Vaupes, cachivera de Yurupari, 400 m, 
Garcia- Barriga 14948 (COL). 
PERU. Without exact locality: Hobb s.n. (BM). 

BRAZIL. Without exact locality: Burchell 3168 (P); Glaziou 1646 (C); 2241 (C, type of S. brevipes Fee 
and S.feei, Hieron.), 3543 (C), 4492 (C, P), 4495 (C, P), 4496 (C), 4498 (C, P), 7038 (C), 7354 (C), 9313 
(C), 13369 (P); Richardson s.n. (BM). Amazonas: near Panure, Rio Uaupes, Spruce 2532 (B, type of S. 
deltoides A.Br.; BM, CGE; K, type of S. trifurcata Baker); Tutica, Uaupes, Luetzelburg 23710 (M). 
Espirito Santo: Rambo 42647 T (BM); Cachoeiro do Itapemim, vargem Alta-Guiomar, Brade 19346 
(BM); Santa Barbara do Caparao, Chase 10075 1/2 (BM). Minas Gerais: Ouro Fino, Hoehne 19459 (S); 
Serra de Antonio Perura, near Ouro Preto, Silveira 171 (P); Christina, Luederwaldt 20999 (BM). 
Parana: Serra do Mar, Cadeodo, Dusen 654a (C); Wielewski 72a (NY), 72 (NY). Santa Catarina: Lagos, 
Spannagel 85 A (NY), 8 5 (BM, NY); Joinville, Muller 1 34 (BM). Rio Grande do Sul: Jurgens L 1 (BM); 
Biopitrade, Santa Cruz, Stier L 25 (NY); Dutra 227 (BM); Apiahy, Glaziou 160 (P); Vila Manresa, 
Rambo 42719 (BM); Rio Pardo, Jurgens L 5 (BM); Taquara, Dutra 668 (BM); Pareci near Montenegro, 
Rambo 49235 (BM); ad montem Ferrabraz near Hamburg, Rambo 43205 (BM); Sao Leopolodo, Dutra 
75 (BM), 1 607 (BM), 1613 (BM); near Belem Novo, 20 km S of Porto Alegre, Beetle 1 592 (BM). Rio de 
Janeiro. Rudio s.n. (B, BM); Glaziou 13365 (B, C); Raddi s.n. (BM); Gardner 67 (BM, K. P), 66 (BM, 
P); Mosen 55 (P), 2022 (P); Brade s.n. (BM); Tijuca, Lutz 1053 (BM); Yucca, Corcovado, Rose & 
Russell 20X99 (NY), 20186 (NY); Murrill s.n. (NY); Tijuca-Bico, Alston & Lutz 307 (BM); Organ mts, 
Rose & Russell 20750 (NY); St Antonio de Finbe, 600 m, Brade 1 1663 (BM); Itatiaia, Hoehne 20996 
(BM); Luederwaldt s.n. (S); Brade 10270 (BM), 10051 (BM); 1000 m, Brade 17189 (BM); 800 m, Brade 
18850 (BM); Monte Serrat, near Itatiaia, Rose & Russell 20441 (NY); Hoehne 24874 (S); Serra do 
Itatiaia, 800 m, Brade 6507 (BM); Itatiaia, estrada Maromba, Bandeira s.n. (BM); Teresopolis, 
cachoeira Teroz, 900-1000 m, Lutz 2081 (BM); Theresopolis, Fazenda Cumary a Cachoeira Feroz, 
810-840 m, Lutz 2107 (BM). Santa Catarina: MacRae s.n. (BM): Schreiner s.n. (BM). Sao Paulo: 
Jundiahy, Puiggari 1 1 (S); Jose Menino, Santos, Gehrt 33427 (BM); Bocaina, Loefgren s.n. (SP); Morro 
do Bahu, Loefgren & Edwall s.n. (SP); Nabraya, Dusen 2148 (P); Santos, Mosen 3125 (C); Jaragua, 
Brade 5137 (NY); Morro de Pedras, Brade 7690 (BM, NY); Serra da Cantareira, Tamandare & Brade 
6598 (NY); Puizzari, Schwacke 2384 (BM); Cantareira, Hoehne 21027 (BM); Morro das Pedras, 
Iguape, Brade 15719 (BM); Campos do Jordao, Leite 3899 (BM); Campo Grande, Serra do Mar, 800 m, 
Brade 6599 (BM). 

Geographical range: Tobago, Trinidad and Guyana, west to Colombia and south to northern 
Brazil; also Argentina, Uruguay and Paraguay. 

Notes: S. muscosa is a variable and possibly mixed species that would merit further 
investigation. Specimens from the Guyanas have been segregated as S. deltoides A. Braun; 
they have lateral leaves which are more triangular, but are otherwise similar. Alston has 
these segregated as a distinct species but in our opinion the difference is within the range of S. 
muscosa. Selaginella deltoides A. Braun was based on Spruce 2532 and 2535; isotype 
material exists in BM and K, but all the material at Berlin belongs to the species 
subsequently described by Baker as S. trifurcata which we believe to be synonymous with S. 
muscosa. Spruce 2535 of other herbaria is S. dendricola Jenman, i.e. S. deltoides sensu 
Baker. A specimen of Spruce 2328, which is S. cordifolia (Desv.) Spring, was subsequently 



SELAG1NELLA IN TROPICAL SOUTH AMERICA 299 

added to the Berlin sheet by A. Braun. S. crassinervia and S. polysperma were reduced to S. 
brasiliensis by A. Braun (Annls Sci. nat. (Bot.) V, 3 : 290; 1865), and S. beyrichii (in Index 
Sem. Hort. bot., Berl. Appendix 1857 : 16; 1857). Hieronymus confused this species with S. 
humilis Jenman, which is S. cladorrhizans (No. 43). S. muscosa may also be confused with 
S. cordifolia (Desv.) Spring but that species has long ciliate lateral leaves and distinctly 
auricled median leaves. S. potaroensis (No. 87) is another similar species, but is 
distinguished by small unequal-sided median leaves and broad, short platystachyous strobili. 
S. jlagellata (No. 41) normally differs by its flagelliform apices and long, narrow, 
platystachyous strobili. 

Lycopodium brasiliense Raddi was the first epithet to be given to this species, but two 
years later Desvaux (Mem Soc. linn. Paris 6 : 190; 1827) described another species by the 
same name (see S. jlexuosa, No. 67); that species was combined in Selaginella in 1838 thus 
making the transfer of Raddi's epithet by A. Braun in 1 865 illegitimate. 

89. Selaginella dendricola Jenman in Gdnrs' Chron. Ill, 2 : 99 (1887); Baker, Fern Allies: 
70 (1887); Jenman, Ferns Brit. W. Indies : 392 (1909); Alston in Reprium Spec. nov. Regni 
veg. 45 : 315 (1936); Knox in Trans, bot. Soc. Edinb. 35 : 264 (1950). Type from Guyana, 
Essequibo: Macouria River, Jenman 2323 (NY, holotype; BM, K, isotypes). 

Selaginella deltoides sensu Baker, Fern Allies : 72 (1887), non A. Braun (1865). 

Specimens seen: 

FRENCH GUIANA. Guya: Devil's Mts, La Comte R., Boulanger Creek, on rocks near the waterfall, 

Leprieur 24$ (BM,P) : 

SURINAM. Marowijne : Nassau Mts, Lanjouw & Lindemann 2628 (BM). 

GUYANA. Essequibo: Type as above; Mt Roraima, Warrington, Burras, Woodhams & Edwards s.n. 

(BM, K); Moraballi Creek, Richards 19 (K). 

COLOMBIA. Vaupes: Cachivera Muriti, Rio Kanararii, 250 m, Schultes & Cabrera 14410b (US); 

Circasia, 240 m, Schultes & Cabrera 19684 (US). 

BRAZIL. Amazonas: Panure, Rio Uapes, Spruce 2535 (BM, CGE, K, US); Cachoeira Cunati, R. Xie, 

Rio Negro, Schultes & Lopez 92 1 9 (BM). 

Geographical range: Colombia to northern Brazil. 

Notes: The elongate stem with small distant leaves, alternating with stems with larger closer 
leaves, is quite distinctive but not shown by all specimens. From S. producta (No. 100) this 
species may be distinguished by its more slender, flexuose, stem, with short filiform 
rhizophores, thinner texture, stouter strobili, and narrower, proportionately longer, median 
leaves. S. seemannii (No. 63) has a similar texture to S. dendricola but the stem is stout, not 
flexuose, and more regularly pinnate, the lateral leaves are conspicuously costate, and the 
median leaves are oblique at the base and shortly auriculate on the outer side. Baker applied 
S. deltoides A. Braun to Spruce 2535 as represented at Kew and most other herbaria, but 
Braun's material (in B) is not this plant (see No. 88). 

90. Selaginella tenuissima Fee in Crypt. Vase. Bres. 2 : 98 (1873). Type from Brazil: Rio de 
Janeiro, Glaziou 4499 (C). 

Selaginella cunninghami Baker in J. Bot., Lond. 22 : 1 10 (1884). Syntypes from Brazil. 
Selaginella papagaiensis Silveira in Bolm Comm. geogr. geol. Minas Geraes 5 : 125 (1898). 

Type from Brazil. 
Selaginella serpens sensu Fee, op cit. : 99 ( 1 873), non (Desv.) Spring ( 1 843). 

Specimens seen: 

BRAZIL. Without exact locality: Glaziou 52 1 7 (BM, syntype of S. cunninghami Baker). Minas Gerais: 
Papagaio, Silveira 154 (BM, type of S. papagaiensis A. Silveira): Ouro Preto, Hoehne 20993 (S): Passa 
Quatro Itaguare, 2000 m, Brade & Araujo 19040 (BM): Serra do Caparao, Rio Domingos, 1900 m, 
Brade 17108 (BM). Rio Grande do Sul: Porto Alegre, Lindman A517 (BM). Rio Janeiro: type, as 
above: Glaziou 4486 (BM, C, K, L), 4485 (C, K); Cunningham s.n. (K, syntype of 5. cunninghami 
Baker): Itatiaia, Hemmendorff'515 (BM): Terezopolis, Brade 9662 (BM); St Antonio de Finbe, Brade & 



300 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Luis 11666 (BM). Santa Catarina: in einem Erdloche des Laranjiera-Gebirges, Vie 131 (BM). Sao 
Paulo: Itatiaia, Hoehne 20996 p.p. (BM), Brade 6508 (NY); Campos do Jordao, 1600 m, Leite 3673 
(BM). 

Geographical range: Endemic to Brazil. 

Notes: This is, perhaps, only a slender state of S. muscosa Spring. 

91. Selaginella falcata (Beauv.) Spring in Bull. Acad. r. Belg. 10:225 (1843), p.p. excl. 
planta borbonica in Mem. Acad. r. Sci. Lett. Belg. 24 : 181 (1850); p.p. nomen tantum; 
Alston in Dansk hot. Ark. 7:200 (1932). Type from French Guiana, without precise 
locality, Chastelein s.n. (OXF). 

Plante muscosa et plumosa guayanensis perelegans major Breyn., Exot. Plant. : 194 (1678); 

Ray, Hist. Plant. 1: 119(1686). 
Muscus Americanus, denticulatus, major Tourn., Lnst. Rei Herb. : 556 (1700), based on 

Breynius. 
Lycopodioides dichostomum taxiforme Dillen., Hist. Muse. : 471, t.66 (1742), copied from 

Breynius. 
Stachygynandrum falcatum Beauv., Prod. Aethog.: 106 (1805), based on Dillen., t.66, p.p. 

excl. pi. maurit. etborbon. 
Lycop odium falcatum (Beauv.) Desv. in Poiret, Encycl. Meth. Suppl. 3 : 540 (181 3 — 4), p.p. 

excl. pi. maurit. et borbon., in Mem. Soc. linn. Paris 6: 188 (1827), p.p. quoad syn. 

Schkuhr. 
Lycop odium plumosum sensu Schkuhr, Krypt. Gewachse 1 : 167 (1809), non L. (1753). 
Selaginella breynii Spring in Martius, Fl. Bras. I (2) : 1 2 1 ( 1 84), p.p. quoad syn. Breynii; sensu 

Schomburgk Reise Brit. Guian.3: 880 (1848), non Spring (1840). 
Selaginella guianensis Spring in Bull. Acad. r. Belg. 10: 143 (1843). Type from French 

Guiana. 
Selaginella guayanensis Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 134 (1850); Baker, Fern 

Allies : 56 (1887). Type as above. 

Specimens seen: 

FRENCH GUIANA. Without precise locality: type, as above; Leprieur s.n. (P, type of S. guianensis 

Spring); R. Oyapok, Leprieur 1 6 1 (BM), 1 62 (BM). Guya: banks of R La Comte, Leprieur 1 80 (K). 

Geographical range: Guyana ? and French Guiana. 

Notes: This species has been much confused. The plant figured by Breynius came from 
French Guiana and there is a scrap at Oxford, collected by Chastelein, which is presumably 
the type collection. Dillenius seems to have copied the figure of Breynius, and Schkuhr's 
plate is from a specimen in Herb. Breynius (BM). 

Palisot de Beauvois (I.e.: 106) based his species upon the plate of Dillenius, without 
description, but cited the 4 Ile-de-France [Mauritius] et de Bourbon [Reunion]' as localities 
on p. 113. The plant from Mauritius was S. concinna Sw. Though Spring took his name from 
Breynius, the description was taken from the plant of Martius, now at Munchen. 
Schomburgk (1 848) records it along the Essequibo and Barima in Guyana. 

92. Selaginella densifolia Spruce ex Hook. Second Cent. Ferns : t.85 (1861); Baker in J. 
Bot., Lond. 21 : 242 (1883), Fern Allies : 55 (1887). Type from Colombia, Vichada: Cerro 
de Mono, between San Fernando and Maypures, Spruce 3089 (K, holotype; BM, CGE, 
LG, isotypes). 

Selaginella orinocensis Maury in J. Bot., Paris 3 : 1 30 (1 889). Type from Colombia. 

Specimens seen: 

COLOMBIA. Vicharda: type as above; loc. idem., Gaillard 201 (P; type of S. orinocensis 

Maury). 

Geographical range: Confined to the Cerrode Mono, Colombia. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 301 

Notes: Both Hooker and Baker cited Spruce 3809 in mistake for 3089, and Cerro de Morro 
for Cerro de Mono. The type of S. orinocensis seems to have been collected in precisely the 
same spot as Spruce's type, but the stems are broader and the leaves slightly acuminate, 
showing some characterstics of S. kochii (No. 93). Typical S. densifolia has the lateral leaves 
tightly curled around the stem, especially on the primary branches, when dry, and in the 
young shoot they are dense and tend to curl upwards in a characteristic manner. 

The Spruce reference (Notes of a Botanist in the Amazonian Andes 1 : 454; 1908) to a 
'Selaginella bearing some resemblance to a closely cut box-edging' was most likely to this 
species. 

93. Selaginella kochii Hieron. in Koch-Griinberg, Reisen in Nordwest-Brasilien 2:361 
(1910), in Reprium Spec. nov. Regni veg. 8 : 151 (1910); Knox in Trans, hot. Soc. Edinb. 
35 : 281 (1950). Type from Brazil, Amazonas: Cachoeira Boredao, Rio Negro, Koch 21 
p.p. (B, holotype; BM, isotype). 

Selaginella densifolia sensu Alston in Pulle, Fl. Surin. 1 : 1 70 (1938), non Spruce. 

Specimens seen: 

SURINAM. Nickeri: Upper Sipaliwini, camp IX, Rombouts 326 (BM). 

VENEZUELA. Amazonas: Piedra de Cocui, Rio Negro, Schultes & Lopez 9430 (BM). 

COLOMBIA. Caqueta: Florencia, Perez Arbelaez 642 (BM, US); Solano, 8 km SE of Tres Esquinas, on 

Rio Caqueta below mouth of Rio Orteguaza, 200 m, Little 9713 (COL), 9545 (COL). Meta: near 

Villavicencio, 500 m, Perez Arbelaez 182 (BM); Alston 7538 (BM). Valle del Cauca: El Silencio, 

Yanaconas, 1900—2200 m, Killip & Garcia 33746 (COL). Vaupes : Rio Kuduyari, Cerro Yapoboda, 

450 m, Schultes & Cabrera 14172 (COL); Cerro Yapoboda, 45 m, Schultes & Cabrera 14174 (US); 

Cerro Isibukuri, 250-700 m, Schultes & Cabrera 13316 (US, COL). 

BRAZIL. Amazonas: type, as above; Serra de Sao Gabriel, Rio Negro, Schultes & Lopez 9176 (BM); 

Serra Curicuriari, Schultes & Lopez 9827 (BM); Serra Wabeesee, Rio Uaupes, Schultes & Pires 9134 

(BM). 

Geographical range: Amazon basin from Surinam to Colombia. 

Notes: The Rombouts specimen from the Upper Sipaliwini is smaller but more like this 
species than S. densifolia (No. 92). 

94. Selaginella truncata Karsten ex A. Braun, Index Sem. Hort. hot., Berl. Appendix 
1857 : 15 (1857), in Annls Sci. nat. (Bot.) IV, 13 : 65 (1860), op. cit. V, 3 : 273 (1865); 
Baker, Fern Allies : 53 (1887); Knox in Trans, bot. Soc. Edinb. 35 : 281 (1950). Syntypes 
from Colombia, Cundinamarca: Bogota, Andes of New Grenada, Karsten s.n. (B, BM); 
Susumuco, 1000 m, Triana 696 (BM); 238 (NY). 

Selaginella weberbaueri Hieron. ex Knox in Trans, bot. Soc. Edinb. 35 : 282 (1950). 

Specimens seen: 

VENEZUELA. Without exact locality: Burro, S. Esteban, Goebel 1889 (P). 

COLOMBIA. Boyaca: Labranzegrande, 1150 m, Guevara Amortegui 345 (BM, US). Cundinamarca: 

syntypes as above; Bogota, Triana s.n. (US). Meta: near Villavicencio, 5-600 m, Perez Arbelaez 182 

[bis] (BM, US); Haught 2524 (BM); Andre 807 (NY): Alston 7612 (BM): Los Llanos, Villavicencio, 

hacia El Parrao, 500 m, Cuatrecasas 4601 (COL); Los Llanos del Rio Guatiquia, near Villavicencio, 

350 m, Molina & Barklev 18M.055 (COL, UC); Llanos Orientales, Santa Isabel, between Restrepo and 

Cumaral, 450 m, Garcia- Barriga 5010 (AAU). 

PERU. Loreto: Mt Guayrapurima, Tarapot, Spruce 4024 (BM, CGE, NY). 

BOLIVIA. La Paz: San Antonio near Mapiri 850 m, Buchtien 1070 (BM, US): Buchtien s.n. 

(Rosenstock exsicc. 95: BM): San Carlos 750 m, Buchtien 54 (US). 

Geographical range: Colombia and Venezuela to Peru and Bolivia. 

Notes: Incomplete specimens may be confused with S. cruciformis (No. 53), which has 
thicker, less flexuose, stems, more oblong median leaves, and lateral instead of terminal 



302 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

strobili. S. breynii (No. 97) has a similar habit, but the median leaves are aristate. S. 
applanata (No. 98) is a smaller plant with the median leaves always ciliate and aristate. 

95. Selaginella homaliae A. Braun in Annls Sci. nat. (Bot.) V, 3 : 274 (1865); Baker, Fern 
Allies : 53 (1887); Alston in Reprium Spec. nov. Regni veg. 45 : 315 (1936). Syntypes from 
Brazil, Amazonas: Cataracts of Jaruma and Panure, Upper Amazon, Spruce 2534 (BM, 
CGE, K); Spruce 2941 (K); Cachoeira da Tarumu, Traill 1437 (K). 

Specimens seen: 

COLOMBIA. Vaupes: Rio Vaupes, Circasia, 240 m, Schultes & Cabrera 19695 (US, COL). 

BRAZIL. Amazonas: syntypes as above. 

Geographical range: Colombia and northern Brazil. 

Notes: This species closely resembles S. applanata (No. 98) in habit, but the median leaves 
are aristate on that species, and their apices are directed forwards instead of overlapping the 
stem. 

96. Selaginella calosticha Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 206 (1850); Baker, 
Fern Allies : 55 (1887); Knox in Trans, bot. Soc. Edinb. 35:280 (1950). Type from 
Venezuela, Federal District: Caracas, 1 500 m, Funck & Schlim 3321 (K). 

Specimens seen: 

VENEZUELA. Federal District: Type as above; Caracas, Funck & Schlim 3320 (BM); Galipan, 

1200 m, Funck & Schlim 335 (BM, C); 3352 (W). 

PERU. San Martin: Tingo Maria, 625-1 100 m, Allard 20821 (BM), 21206 (BM). 

Geographical range: Venezuela and Peru. 

Notes: The microspores have capitate papillae, but the larger size and closer leaves appear to 
distinguish this species from S. tarapotensis (No. 99), which is otherwise very similar. 

97. Selaginella breynii Spring in Martius, Fl. Bras. 1 (2) : 121 (1840), p.p. excl. syn. Breyn.; 
in Mem. Acad. r. Sci. lett. Belg. 24 : 119 (1850), p.p. quoad Schomburgk 982 et spec. 
Martius; Baker, Fern Allies : 54 (1887), p.p. excl. syns Schkuhr et Dillen.; Knox in Trans, 
bot. Soc. Edinb. 35 : 279 (1950). Type from Brazil, Martius s.n. (M). 

Selaginella panamensis Linden, Cat. 10 : 22 (1855), nom. nud. 
Lycopodium concinnum sensu J. Smith in J. Bot., Lond. 1 : 203 (1 842), non Sw. 
Selaginella concinna sensu Schomb., Reise in Guiana 3:880, 1042 (1848), non. (Sw.) 
Spring (1838) nee Baker (1891). 

Specimens seen: 

COLOMBIA. Santander: near Barracan Bermeja, 100-500 m, Haught 1339 (BM); km 6 between 

Puerto Wilches and Puerto Santos, 100-1 15 m, Killip & Smith 14872 (NY). Meta: Pico Renjifo, Serra 

de la Macarena, Philipson, Idrobo & Jaramillo 2 1 62 (BM); Idrobo & Schultes 1151 (BM). 

BRAZIL. Without precise locality: type as above; Glaziou 14414 (BM); Serra de Jurutz, Traill 1432 

(K). Amapa: Oiapoque, Black 8275 (BM); Igarape Pontanarre Rio Oiapoque, Froes 26005 (BM). 

Amazonas: near Manaos, Spruce s.n. (BM, C); Cararaucu, Traill 1434 (K); Conoceicao, Rio Negro, 

Traill 1433 (K); forest on Rio Marnelhos, R. Madeira, Barra, Rio Negro, Spruce s.n. (BM, CGE); 

Cachveira Tamura, Manaos, Occhioni 69 (BM); Igarape do Gnedes, Kuhlmann 998 (BM); Rio Apahu, 

Hubner 56 (BM). Bahia: Sao Bento, Luetzelburg 304 (BM). Para: Rio Padauiri, Schomburgk 982 

(BM); Rio Aripecuru, Spruce s.n. (BM, CGE); Santarem, Spruce s.n. (CGE). 

Geographical range: Brazil and Colombia. 

Notes: The plant from Rio Aripecuru is smaller, as are the specimens from Colombia. The 
record for British Guiana (sub. S. concinna) appears to be based on the Schomburgks 
Brazilian specimen. Spring's record for the Orinoco was based on the same specimen, and his 
Chilean record refers to a specimen of a Philippine species wrongly localised in Haenke's 
collection. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 303 

98. Selaginella applanata A. Braun in Annls Sci. nat. (Bot.) V, 3 : 274 (1865). Type from 
Peru, Puno: near San Gaven, Lechler 2405 (K; BM, fragment). 

Selaginella jungermannioides sensu Baker, Fern Allies : 53 (1887), p.p. quoad pi. peruviana, 
non (Gaudich.) Spring ( 1 843) nee Zoller ( 1854). 

Specimens seen: 

COLOMBIA. Caqueta: Morelia, 300 m, v. Sneidern A 1291 (S). 

PERU. Puno: type as above. 

Geographical range: Colombia and Peru. 

99. Selaginella tarapotensis Baker in J. Bot., Lond. 21 : 98 (1883), Fern Allies : 44 (1887). 
Type from Peru, Loreto: Mt Guayrapurima, near Tarapoto, Spruce 4625 (K, holotype; 
BM, CGE, US, isotypes). 

Specimens seen: 

COLOMBIA. Caldas: La Selva, Pueblo Rico, 1500 m, v. Sneidern 5517 (S). Meta: Cordillera 
Macarena, 1300-1900 m, Schultes 12120 (BM), 12123 (BM, UC), 12125 (BM, UC); Idrobo & Schultes 
1054 (BM, COL), 1055 (BM, COL), 1077 (BM, COL), 1091 (BM). Narino: below Paramo, 1000 m, 
Alston %552 {KM). 

ECUADOR. Santiago-Zamora: Rio Itzintza, Cordillera Cutucu, 1050-1 110 m, Camp E 1221 (BM). 
PERU. Loreto: type as above. San Martin: near Tingo Maria, 846M, Allard 21390 (BM). Junin: 
Schunke Hacienda, above San Ramon, 1300-1700 m, Schunke A 246 (BM, US); Chanchamayo 
Valley, Schunke 195 (BM). Cuzco: Sapansachayoc, Paucartambo, 950 m, Vargas 7350 (BM). 
BOLIVIA. La Paz: Santa Barbara, 1650 m, Williams 1401 (NY, US); Ticunhuaya, 1500 m, Tate 1077 
(US); Hacienda Simaco, on way to Tipuani, Larecaja, 1400 m, Buchtien 5276, (BM, NY, US), 5283 
(BM, NY, US); Polo-Polo, near Coroico, N Yungas, Buchtien s.n. (US); San Antonio, Chapiri, 850 m, 
Buchtien 1045 (US), 1071 (BM, US); San Carlos, Mapiri, 850 m, Buchtien 55 (US), 56 (NY, US), 58 
(NY), 59 (NY), 1078 (BM, US); above Mapiri, 1200 m, Williams 1406 (NY, US); Mapiri, 750 m, 
Rusby 461 A (NY). 

Geographical range: Mexico to Bolivia. 

100. Selaginella producta Baker in J. Bot., Lond. 21 : 243 (1883), p.p., Fern Allies: 56 
(1887), p.p. excl. pi. Lindbergi; Knox in Trans, bot. Soc. Edinb. 35 : 262 (1950). Types 
from Guyana: Appun 196, 198; Drake s.n. (K); and Brazil, Amazonas: between Barcellos 
and San Gabriel, Spruce 2043 (BM, holotype; CGE, isotype); Minas Gerais, Lindberg s.n. 
(K). 

Selaginella cayennensis Baker, Fern Allies : 44 (1887). Type from French Guiana. 
Selaginella tobagensis Hieron. in Symb. Antill. 3 : 524 (1903). Type from Tobago. 
Selaginella tarumensis Hieron. ex Bonap., Notes Pterid. 1 : 1 80 (19 1 5), nom. nud. 

Specimens seen: 

SURINAM. Saramacca: Nassau, Lanjouw & Lindeman 2579 (BM), 2792 (BM); between Pakka-pakka 
and Ebbatop, Florschutz 1 350 (BM); Tafelberg, Maguire 243 1 6 (BM). 

FRENCH GUIANA. Guyane: Bradel Falls, R. le Comte, Leprieur 157 (BM, K, P; type of S. 
cayennensis Baker); covering the rocks at the Bradel, Leprieur 242 (P); Cacao Creek, R. le Comte, 
Leprieur 1 56 (P). 

GUYANA. Without exact locality: syntypes as above; between Demerara and Berbice Rivers, De la 
Cruz 1609 (NY); region of Mt Raywa, Jenman s.n. (NY); near Mt Arisaru, Guppy 29 (BM), 38 (BM). 
Berbice: Sierra Acarai Region, Bay forest, 5 miles East of Onoro, Guppy 243 (BM); Sierra Acarai 
Region, Upper Chodiker, Guppy 461 (BM). Demerara: Great Falls, Demerara R, Jenman s.n. (NY); 
Beccari s.n. (FI); Demara R, Jenman s.n. (NY), 4212 (BM); Werebaru Creek to Baboon camp, Beccari 
s.n. (FI); Baboon camp, Demerara R, Beccari s.n. (FI); Canister Falls, Waini R, Beckett s.n. (BM). 
Essequibo: Sebai Creek, Kaituma R, Fanshawe 2417 (BM); Kanuku Mts, Takubu R, A. Smith 3293 
(BM); Macouria creek, Essequibo R, Jenman s.n. (NY); 2325 (BM); Bartica to Potaro Rd, near 14 m to 
post, Sandwith 1 139 (BM, K); Moraballi Creek, near Bartica, Richards 268 (BM, K) 196 (BM, K); near 
Dadanawa, Upper Rupununi, De la Cruz 1810 (NY); Junction of Mazuruni andCuyuni rivers Graham 
315 (NY); Mazuruni R, near First Falls, Sandwith 1 168 (BM, K); Upper Mazuruni R, De La Cruz 



304 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

2188 (NY, K); Eagle Mt, Potaro R. Jenman s.n. (BM, NY); Tumatumari, Potaro R, Gleason 63 (NY); 

Tiger Creek, Tumatumari, Tutin 548 (BM); Potaro R, Jenman s.n. (NY); Kaieteur Gorge, Tutin 523 

(BM); Sandwith 1477 (BM, K); between Tukeit and Kaieteur Plateau, Maguire & Fanshawe 23089 

(BM),Gwppvl62(BM). 

VENEZUELA. Delta Amacuro: Lower Orinoco, Rusby & Squires 438 (BM, NY), 439 (BM, NY); 

Imataca Mts, Thompson 18 (BM). 

COLOMBIA. Amazonas: Randal Yayacopi, Rio Apaporis, 240 m, Schultes & Cabrera 16216 (US); 

Cerra de la Cente Chiquita, Schultes & Cabrera 16497 (BM, US). Narino: Buenaventura, Andre 252 

(NY). Santander: near Barranca Bermeja, 100-500 m, H aught 1338 (BM). 

PERU. Loreto: Iquitos, 100 m, Killip & Smith 27322 (BM, NY); Mishuyacu, 100 m, Klug 198 (BM, 

NY, US). 

BRAZIL. Without exact locality: Glaziou 10212 (C), 10215 (C). Amazonas: syntype as above; without 

further locality, Schwacke s.n. (BM); Taruma rapids near Barra, Spruce 473 1 (BM, CGE, US; type of S. 

tarumensis Hieron.); Occhioni 20220 (BM). Goias: Corrego Alogado, c. 30 km, SW of Brasilia, 1000 m, 

Irwin, Grear, Souza & Reis dos Santos 14056 (BM, NY); Corrego Itaquera, c. 30 km N of Formosa, 

850 m, Irwin, Souza, Grear & Reis dos Santos 15573 (BM, NY). Parana: Palmeira, Freitas 55 (BM, 

NY). 

Geographical range: Tobago and Trinidad to French Guiana, west to Colombia and south to 
Brasilia and northern Peru. 

Notes: S. revoluta (No. 102) is superficially similar but easily separated by the long cilia at 
the base of the lateral leaves. These lateral leaves are often inrolled and are usually smaller, 
narrower and more acute than the median leaves. S. seemannii (No. 63) is similar in habit, 
but the lateral leaves are thinner with a distinct costa, and the median leaves gradually 
acuminate rather than abruptly aristate, with oblique auriculate bases. S. cayennensis Baker 
is a small form, and is matched by Tutin 548 from wet rocks in Guyana; it is within the range 
of the species. 

101. Selaginella macrostachya (Spring) Spring in Bull. Acad. r. Belg. 10 : 144 (1843). Type 
from Brazil, without exact locality, Sellow s.n. (K). 

Selaginella flexuosa subsp. macrostachya Spring in Martius, Fl. Bras. 1 (2): 123 (1840). 

Type as above. 
Selaginella gardneri Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 134 (1850). Type from 

Brazil. 
Selaginella ericoides Fee, Crypt. Vase. Bres. : 228 (1869); Hieron. in Hedwigia 43:38 

(1903). Type from Brazil. 
Selaginella geminata Fee, Crypt. Vase, Bres., Suppl. : 100 (1873). Type from Brazil. 
Selaginella lindbergii Baker in J. Bot., Lond. 21 : 99 (1 883). Type from Brazil. 
Selaginella henriqueana Silveira in Bolm Comm. geogr. geol. Minas Geraes 5:123 

(1898). Type from Brazil. 

Specimens seen: 

BRAZIL. Without exact locality: type as above; Glaziou 2243 (BR, type of S. ericoides Fee), 4500 (C). 
Minas Gerais: Lindberg s.n. (K, type of S. lindbergii Baker); near Chora, Silveira 163 (BM). Rio de 
Janeiro: Glaziou 4484 (C, K, type of S. geminata Fee); Alto Macahe, Glaziou 7281 (BM, C, P), 7282 
(BM, P); Serra das Orgaos, Gardner 5958 (K, type of S. gardneri); Serra dos Orgaos Corrigo Berjaflos, 
1300 m, Brade 16585 (BM); Teresopolis, Brade 9444 (BM); Teresopolis, Cachoeira Teroz, 
900-1000 m, Lutz 2080 (BM). Santa Catarina: Fachinal Biguassu, Rambo 50372 (BM); Brusque, 50 m, 
Reitz 3144 (BM); Morro Queimadas, Spannagel 212 (NY); Alto da Serra, Bauer 81 (NY). Sao Paulo: 
Alto da Serra, Brade 5851 (BM); Hoehne 4735 (BM), 28172 (S); Gehrt s.n. (BM); Edwall s.n. (BM); 
Luederwaldt 21034 (S); Parque Cazuru, Luederwaldt 1657 (BM); Santos, Mosen 3758 (C); Serra do 
Itatins, Iguape, 800 m, Brade 8627 (BM). 

Geographical range: Confined to southern Brazil. 

102. Selaginella revoluta Baker in J. Bot., Lond. 21 : 141 (1883), Fern Allies : 46 (1887); 
Alston in Pulle, Fl. Surinam 1 : 169 (1938); Knox in Trans, bot. Soc. Edinb. 35 : 260 



SELAGINELLA IN TROPICAL SOUTH AMERICA 305 

(1950). Type from Venezuela, Amazonas: near Maupures, Spruce 3621 (K, holotype; BM, 

CGE, US, isotypes). 
Selaginella platybasis Baker in J. Bot., Lond. 21 : 242 (1883), Fern Allies : 54 (1887); Knox 

in Trans, bot. Soc. Edinb. 35 : 262 (1 50). Type from Brazil. 
Selaginella demissa Christ in Bull. Herb. Boissier II, 1 : 75 (1901). Type from Peru. 

Specimens seen: 

FRENCH GUIANA. Without precise locality, Leprieur s.n. (NY, P). Inini: Upper Oyapok, Leprieur A 

(NY, P); Naneteri, Upper Oyapok, Leprieur 169 (P). 

SURINAM. Nickerie: Nickerie R., Tulleken 311 (L, U). Saramacca: Wilhelminagebergte, Stahel 335 

(U),482(U). 

GUYANA. Essequibo: Eagle Mt, Potaro R, Jenman s.n. (BM, NY); Tiger Creek, Tametumari, Tutin 

544 (BM); Garroway stream Potaro River gorge, Maguire & Fanshawe 2291 \ (BM). 

VENEZUELA. Amazonas: type, as above; Rios Pacimoni-Yatua, Casiquiare, on Piedra Aravicava, 

1 10—200 m, Maguire, Wurdack & Bunting 37458 (BM); Bolivar: salto de Para, Medico Caura, 300 m, 

Williams U379 (BM). 

COLOMBIA. Caqueta: Florencia, Perez Arbelaez 668 (BM, US). Meta: Rio Guapaya, Macarena, 

450 m, Philipson, Idrobo & Fernandez 1607 (BM, COL). Narino: Buenaventura, Andre 273 (NY). 

Valle del Cauca: Cali-Buenaventure Highway, 100 m, Haught 5320 (BM, COL). Vaupes: below 

Teresita, Rio Papuri, Schultes & Cabrera 19484 (US); Cano Paca, Rio Piraparana, Schultes & Cabrera 

17595a (BM, US); Soratama, between Rio Pacoa & Rio Kananari, Schultes & Cabrera 13022 (US), 

13751 (US). 

PERU. Loreto: Rio Mayo, near Tarapoto, Spruce 4732 (BM); Mishuyacu, near Iquitos, 100 m, Klug 

1220 (BM, NY, US); Gametanacocha, Rio Mazari, 100-125 m, Schunke 322 (BM, UC); Ucayali, 

Ruber s.n. (P, type of S. demissa). San Martin: near Tingo Maria, 625-1 100 m, Allard 20415 (BM), 

20485 (BM), 22396 (BM). 

BRAZIL. Amazonas: San Gabriel de Cachoeira, Spruce 2195 (BM, CGE); Panure on Rio Uapes, 

Spruce 2502 (BM, type of 5. platybasis); summit of Serra Curicuriari, Schultes & Lopez 9839 (BM). 

Geographical range: French Guiana to Panama south to 10°S latitude. 

Notes: This species has the habit of S. producta (No. 100) but is separated by its ciliate, 
tapering, and more acute, lateral leaves. The leaves are sometimes slightly pubescent on the 
upper surface near the margins (e.g. Tutin 544). 

103. Selaginella porelloides (Lam.) Spring in Bull. Acad. r. Belg. 10 : 141 (1843); Hieron. in 

Hedwigia 43 : 298 ( 1 9 1 7). Type from 'Croit dans les Antilles' (P, not seen). 
Lycopodium porelloides Lam., Encycl. 3 : 652 (1791). Type as above. 
Stachygynandrum porelloides (Lam.) Beauv., Prodr. Aethog. : 1 10 (1805). Type as above. 
L. anomalum Hook. & Grev. in Hook., Bot. Misc. 2 : 400 (1831). Type from Guyana. 
S. anomala (Hook. & Grev.) Spring in Bull. Acad. r. Belg. 10 : 232 (1843), in Mem. Acad. r. 

Sci. Lett. Belg. 24 : 247 (1850); Baker, Fern Allies : 122 (1887). Type as above. 
S. platyphylla Baker in J. Bot., Lond. 23 : 294 (1885), Fern Allies : 121 (1887), non Veitch 

( 1 882). Type from Guyana. 
S. platyphylla var. laxa Jenman, Ferns Brit. W. Indies : 389 (1909). Type from Guyana. 
S. jenmani Baker, Fern Allies : 45 (1887), in Gdnrs' Chron. Ill, 2 : 154 (1887). Type from 

Guyana. 

Specimens seen: 

FRENCH GUIANA. Without precise locality: Leprieur s.n. (NY), 168 (BM, P); Oyapock, Leprieur 

s.n. (P); Karouany, Sagot 749 (BM, NY, P). Guyane: stream at base of Mt Matouri, Cayenne Isl., 

Leprieur 165 (K); Maroni, Portal Isl., Sagot 1 123 (K); bank of R Sourou, R la Comte, Leprieur 167 (P); 

near Cayenne, Mille s.n. (P). 

GUYANA. Without precise locality: Jenman 1818 (K), 1819 (K, type of S. platyphylla var. laxa 

Jenman); Mt Raywa, Jenman s.n. (NY). Demerara: Upper Demerara R, Jenman s.n. (BM, K, syntype 

of S. platyphylla Baker; NY): Kara Kara Creek, Demerara R, Jenman s.n. (NY); Demerara, Ankers s.n. 

(K, type of S. anomala Spring). Essequibo: Bartica Grove, Jenman 2322 (BM); Baracaa, Mazaruni, 

Essequibo R, Barkly s.n. (BM); between Demerara and Berbice rivers, De la Cruz 1608 (NY); Kaieteur 

ravine, im Thurn 78 (K, syntype of S. platyphylla Baker); ravines near Kaieteur Fall, Jenman s.n. (NY); 



306 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

ravines near Kaieteur savannah, Jenman 1482 (K); foot of the Kaieteur, Jenman 1480 (BM, K, type of 
S.jenmani Baker; NY); between Tukeit and the Kaieteur savannah, Sandwith 1452 (BM, K); Upper 
Rupununi, near Dadanawa, De la Cruz 1529 (NY); near Bartica, Essequibo R, De la Cruz 1935 (NY); 
forest opposite Bartica, Jenman s.n. (NY); Macouria creek, Essequibo, Jenman s.n. (NY); Sandwith 
1568 (BM, K); H.M.P.S., near Kartabo, Essequibo R, Guppv 1 (BM); Mazaruni Station, Tutin 62 
(BM). 

Geographical range: Confined to the Guianas. 

104. Selaginella exaltata (Kunze) Spring in Bull. Acad. r. Belg. 10 : 234 (1843), in Mem. 
Acad. r. Sci. Lett. Belg. 24 : 145 (1850); Baker, Fern Allies : 93 (1887); Knox in Trans, hot. 
Soc. Edinb. 35 : 270 (1950). Type from Peru, San Martin: ab Uchiza ad Tocoche, Poeppig 
1953 (syntypes ?B, LG; not seen). 

Lycopodium exaltatum Kunze in Linnaea 9 : 8 (1835). Type as above. 
Selaginella strobilifera Christ in Bull. Herb. Boissier II, 1 : 72 (1901), in Hedwigia 45 : 194 
(1906). Type from Peru. 

Specimens seen: 

COLOMBIA. Amazonas: Hervinha, Loreto-Yaco, Black & Schultes 46-258 (BM); Carlo Guacaya, Rio 
Miritiparana, 215 m, Schultes & Cabrera 16568 (US). Antioquia: Vijagual, S of Turbo, 40 m, Haught 
4555 (BM). Choco: Rio Jurado, 100 m, v. Sneidern A 184 (S); Rio Truando, Schott 5 (NY). Narino: 
Barbacoas, Alston 8504 (BM); Pambana, Rio Telembi, 50 m, Ewan 16833 (BM); Costa del Pacifico, 
Herrara, Candelillas 30-100 m, in valley of Rio Mira, Idrobo & Weber 1392 (COL). Putumayo; Puerto 
Ospina, Schultes & Cabrera 18945 (US). 

ECUADOR. Without exact locality; Fraser s.n. (BM); Esmeraldas, Sodiro s.n. (NY). Napo-Pastaza: 
Archedona, Jameson 708 (BM); 650 m, Mexia 7281 (BM); Tena, Asplund 8893 (S); Vera Cruz on Rio 
Indillama, Asplund 19448 (S); Puerto Francisco de Orellana 300 m, Balslev & Madsen 10604 (BM). 
PERU. Junin: between Azupizu and Santa Rosa, 650 m, Killip & Smith 26136 (BM, NY); Puerto 
Bermudez, 375 m, Killip & Smith 26498 (BM, NY). Loreto: Santa Rosa, Rio Huallaga, 135 m, Killip & 
Smith 28776 (BM, NY); between Yurimaguas and Balsapuerto, 135-150 m, Killip & Smith 28088 
(BM, NY); Rio Huallaga, 155-210 m, Williams 4825 (BM); Pebas, Williams 1600 (BM). 
BRAZIL. Without exact locality, Glaziou 10216 (C). Acre; Haradouro do Abunam, Kuhlmann 1722 
(BM). Amazonas; Monte Verde, Rio Purus, Ruber 4555 (BM). 

Geographical range: Panama, south to Peru and western Brazil. 

Notes: Spring (1 850) states that the stems may be up to 18m long. Asplund (on specimen no. 
8893) notes 'scandent to a height of 3 m'; other collectors give lower figures. 

105. Selaginella tomentosa Spring in Mem. Acad. r. Belg. 24 : 231 (1850). Knox in Trans, 
bot. Soc. Edinb. 35 : 290 (1950). Type from Colombia, Cauca: Gorgona Isl., Hinds s.n. 
(K). 

Selaginella geniculata var. tomentosa (Spring) Baker in /. Bot., Lond. 23 : 121 (1885), Fern 

Allies: 105 (1887). Type as above. 
Selaginella ferruminata sensu Seemann, Bot. Herald: 243 (1855), non Spring (1843). Type 

from Peru. 

Specimen seen: 

COLOMBIA. Antioquia: Rio Ampurrumiado, 200 m, Gutierrez & Barkley 17C151 (BM). Cauca: Isle 

of Gorgona, Barclay 902 (BM); Cheesman 26 (BM); type as above. Choco: Bay of Choco, Seemann 

1005 (BM). Vallee del Cauca: bridge over Dagua, near Buenaventura, Alston 8634 (BM). 

ECUADOR. Santiago-Zamora: Cordillera Cutucii, Camp E 1066 (BM). Tungurahua: Volcano 

Tungurahua, Sodiro s.n. (NY). 

Geographical range: Confined to Colombia and Ecuador. 

106. Selaginella conduplicata Spring in Martius, Fl. Bras. 1 (2) : 129 (1840); in Flora, Jena 
21 : 200 (1838), nom. nud., in Mem. Acad. r. Sci. Lett. Belg. 34 : 229 (1850), p.p. quoad pi. 
bras. Type from Brazil, Para, Martius s.n. (M). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 307 

Selaginella stellata Spring in Martius FL Bras. 1 (2): 129 (1840), p.p. non Spring (1838), 
Mem. Acad. r. Sci. Lett. Belg. 24 : 228 (1850); sensu Alston in Reprium Spec. nov. Regni 
veg. 40 : 309 (1936); sensu Pulle, FL Surinam 1 : 163 (1938). Type from Brazil. 

Selaginella calcarata A. Braun in Annls Sci. nat. (Bot.) V, 3 : 305 (1865). Type from Brazil. 

Selaginella parkeri var. stellata (Spring) Baker in J. Bot., Lond. 23 : 120 (1 885), Fern Allies : 
104(1887). Type as above. 

Specimens seen: 

FRENCH GUIANA. Guyane: Cayenne, Leprieur s.n. (NY), 2 (P); Porteau s.n. (K). Inini: Rio Gabaret, 
Oyapok, Leprieur 1 74 (K). 

SURINAM. Without precise locality: Focke 1217 (K, U). Marowijne: Rio Corentyne near 
Poelegoedoe, Versteeg 633 (U). 

COLOMBIA. Amazonas: Cano Guacaya, Rio Miritiparana, 215 m, Schultes & Cabrera 16394 (US). 
Santander: near Barranca Bermeja, Magdalena Valley, between Sogamoso and Colorado Rivers, 
100— 500 m, Haught 1266 (BM). Vaupes: Cerro de Mitu, 250 m, Schultes & Cabrera 13937 (US); 
Cachivera Palito, Rio Kananari, 250 m, Schultes & Cabrera 13177 (BM, US); Yapaboda, Rio 
Kuduyari, 275-305 m, Schultes & Cabrera 20028 (US); Rio Kuduyari, 2 1 5-245 m, Schultes & Cabrera 
20020 (US); Lagunas del Churucho, Rio Apaporis, 280 m, Garcia 13661 (BM); Cachivera de Jirijirimo, 
Rio Apaporis, 250 m, Schultes & Cabrera 12438 (COL, US); Rio Pacoa, tributary Rio Apaporis, 
900 m, Schultes & Cabrera 15200 (COL, US); Soratama, Rio Apaporis, 275 m, Schultes & Cabrera 
15020 (COL, US); Rio Vaupes, Cerro Mitu, Schultes & Cabrera 17827 (COL); Rio Apaporis, 
Soratama, Schultes & Cabrera 16150 (COL). 

PERU. Loreto: Iquitos, Asplund 13939 (S); Mishuyacu, near Iquitos, 100 m, Klug 150 (NY); Iquitos, c. 
100 m, Killip & Smith 26937 (BM), 27030 (BM, NY), 27376 (BM, NY); trail to San Juan, Iquitos, 
Mexia 6496 (BM); Amazon River, near Iquitos, 120 m, Williams 8237 (BM). 

BRAZIL. Without exact locality: Lippald s.n. (BM); Burchell 9436-2 (K), 9269 (K). Amazonas: 
Glaziou s.n. (BM); Manaos, Kuhlmann 980 (BM); Killip & Smith 30101 (BM); KrukoffW\5 (BM); 
Beach Taraina, Manaos, Solomon s.n. (BM); Cerro Dimiti, Rio Negro, Schultes & Lopez 9950 (BM); 
mouth of Rio Xie, Schultes & Lopez 9203 (P); Sao Manoel, Rio Tapajoz, Kuhlmann 67 (BM); Rio 
Taruma, Froes 24860 (BM, IAN); Aurura, Manaos, Luetzelburg 22065 (M); Muyrapenima, Rio 
Negro, Tate 56 (NY); Manaos, Killip & Smith 30101 (BM); Aranja, R Uruara, Traill 1422 (K). Para: 
Thome Assu, Rio Assu, 35 m, Mexia 6003 (BM); Para, Home s.n. (BM); Smith 1 1005 (BM); Spruce 
s.n. (BM, CGE); Ridley, Lea & Ramage 36 (BM); Huber 1265 (BM); L P. G. Smith 284 (BM); 
Prainnha, Rio Pixuna, Black 47-2032 (BM); Tefe, Black 47-1 572 (BM); Tapana, Killip & Smith 30356 
(BM); Cachoeira do Tronco, Rio Cumina, Sampaio 4973 (BM); Utinga, near Belem, Snethlage 1 1 (B); 
Altar do Chao, Traill 1423 (K); Utinga, near Para, Lutz s.n. (BM). 

Geographical range: Guiana and Amazonian Brazil to Peru. 

Notes: 'Powdered leaves used on arms to protect from poisonous brew during preparation by 
witch doctors' (from a note on Schultes & Cabrera 1 5020). 

107. Selaginella asperula Spring in Martius, Fl. Bras. 1 (2): 127 (1840), in Bull. Acad. r. 
Belg. 10:230 (1843); Mem. Acad. r. Sci. Lett. Belg. 24:225 (1850); Baker, Fern 
Allies: 105 (1887); Alston in Reprium Spec. nov. Regni veg. 40:309 (1936): Knox in 
Trans, bot. Soc. Edinb. 35 : 251 (1950). Types from Brazil, Bahia: Barra, Panure: Spruce 
1317 (CGE, not seen). 

Selaginella ovalis Baker in J. Bot, Lond. 21 : 143 (1883), Fern Allies : 48 (1887). Type from 
Brazil. 

Specimens seen: 

VENEZUELA. Amazonas: Cerro Duida, 1 50 m, Stevermark 57742 (BM); 945 m, Tate 890 (BM, NY); 

Esmeraldas, about 100 m, Tate 201 (BM, NY), 327 (NY); Capibara, about 100 m, Holt & Gehringer 

283a (BM, NY); Yerba, 250 m, Molina & Barkley 1 8.v. 1 94 (COL). 

COLOMBIA. Amazonas: Rio Atabapo, 1 km W of Cacagual, Maguire, Wurdack & Bunting 36302 

(BM). Caqueta: Florencia, Perez Arbelaez 621 (BM); Solano, 8 km SE of T res Esquinas, on Rio 

Caqueta below mouth of Rio Orteguaza, 200 m, Little 9714 (COL). Meta: Gorge of R. Guejar, Sierra 

Macarena, 450 m, Philipson 2375 (BM); near Villavicencio, 480 m, Alston 7571 (BM); La Macarena, 

Sabanas de San Juan de Arama, 500 m, Idrobo 2619 (BM); La Macarena (Parte Sur), Rio Guayabero, 



308 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Sabanas de Arenisca, 235-700 m, Barriga & Mejia 17063 (AAU). Vaupes: near Piedra de Cocui, 
Schultes & Lopez 9522 (BM); Mesa la Lindosa, 4-600 m, Idrobo & Schultes 646 (BM, COL), 665, (BM, 
COL); Cerro Yapoboda, Rio Kuduyari, 450 m, Schultes & Cabrera 14216 (US), 14232 (COL, US), 
20066 (US); Puerto Colombia, Rio Guainia, c. 250 m, Schultes, Baker & Cabrera 1794 (COL, US); 
San Felipe, Rio Negro, 185 m, Schultes, Baker & Cabrera 18125 (US); Cerro Isibukuri, Rio Kananari, 
700 m, Schultes & Cabrera 13313 (COL, UC, US); Randal de Yurupari, Schultes & Cabrera 19725 
(US); Savannah Goo-ran-hoo-da, Mesa de Yambi, Rio Karuru, c. 300 m, Schultes & Cabrera 19162 
(US); Cerro Kanenda Rio Kubiyu, 8-900 m, Schultes & Cabrera 18344 (US); Rio Parana Pichuna, 
215 m, Schultes & Cabrera 19912 (US); Cerro E-ree-ee-ko-mee-6-kee, Rio Piraparana, Rio Apaporis, 
Schultes & Cabrera 1 7493 (US); Jinogoje, Rio Apaporis, 215m, Schultes & Cabrera 16589 (US). 
PERU. Without precise locality; Chanchamajo, Schunke s.n. (BM). Junin: La Merced, 760 m, 
Sandeman 5024 (BM); Quimiri Bridge, La Merced, 800—1300 m, Killip & Smith 23833 (BM, NY); 
Colonia Perene, 680 m, Killip & Smith 24942 (BM); bank of Rio Perene, 1260 m, Gascoyne-Cecil 20 
(BM), 670 m, Gascoyne-Cecil 82 (BM); Schunke Hacienda, above San Ramon, 1300-1700 m, Schunke 
A 250 (BM); San Ramon, 900-1300 m, Killip & Smith 24785 (NY). San Martin; Tingo Maria, 
625-1 100 m, Allard 20816 (BM), 20823 (BM), 21647 (BM), 21647a (BM); San Roque, 1350-1500 m, 
L. Williams 14\5(BM). 

BOLIVIA. Tumupasa, 550 m, R. Williams 1396 (BM, NY). 

BRAZIL. Amazonas; Manaos, Spruce s.n. (B, BM, CGE); Solomon s.n. (BM); Schwacke 188 (BM); 
Tate 47 (BM); Killip & Smith 30176 (BM); Campinarana, near Manaos, Krukoff 8015 (BM); Rio 
Mauhes, Traill 1416 (BM, K, type of S. ovalis Baker); Rio Xie, Upper Rio Negro, Schultes & Lopez 
9202 (BM), 9218 (BM); Rio Taruma, Froes 24862 (BM, IAN); Taracua, Rio Uaupes, Schultes & Pires 
9031 (BM); Ipanore, Rio Uaupes, Schultes & Pires 9085 (BM), 9086 (BM); Taracua, Pires 910 (BM). 
Mucajai, Rio Branco, Black & Magalhaes 12943 (BM, IAN); Rio Ayari, Luetzelburg 22505 (M); 
Tunuy, Luetzelburg 22944 (M); Tutica, Uaupes, Luetzelburg 23643 (M); Upper Rio Negro, Weiss & 
Schmidt s.n. (NY). Para: Posto dos Indios Caiabi, Rio Sao Manoel, Pires 386 1 (BM, IAN). 

Geographical range: Amazonian Venezuela and Brazil south to Bolivia. 
Notes: S. ovalis Baker was based on a prostrate, sterile branch of this species. 

108. Selaginella fragilis A. Braun in Annls Sci. nat. (Bot.) V, 3 : 305 (1865), in Mber. K. 
preuss. Akad. Wiss. 1865 : 205 (1866); Alston in Reprium Spec. nov. Regni veg. 40 : 309 
(1938). Type from Brazil, Amazonas: nearPanure, Rio Uapes, Spruce 2533 (BM,CGE). 

Selaginella vanheurckiana Spring in Van Heurck, PL Nov. : 28 (1870). Type as above. 

Selaginella parkeri var. vanheurckiana (Spring) Baker in/. Bot., Lond. 23 : 120 (1885). Type 
as above. 

Selaginella brachylepis Christ in Bull. Herb. Boissier, II, 1 : 74 (1901). Type from Peru. 

Specimens seen: 

VENEZUELA. Amazonas: near Salto de Hua, Serra Imeri, Holt & Blake, 504 (US); Rios Pacimoni- 
Yatua, Casiquiare, 1 1 0—1 30 m, Maguire, Wurdack & Bunting 37469 (BM). 

COLOMBIA. Amazonas: Rio Miritiparana, Cano Guacaya, 215 m, Schultes & Cabrera 15744 (UC). 
Amazonas-Vaupes: Rio Apaporis, Soratama, Schultes & Cabrera 16174 (COL). Antioquia: on 
Quebracha La Puya, east of Turbo, c. 75 m, Haught 4955 (COL). Putumayo: Umbria 325 m, Klug 
1778 (BM). Vaupes: Cerra de Tipiaca, between Mitu and Javarete, Schultes & Cabrera 19303 (US); 
Cerro de Mitu, 250 m, Schultes & Cabrera 13930 (COL, US); below Teresita, Rio Papuri, Schultes & 
Cabrera 19476 (US); Cerro E-ree-ee-ko-mee-6-kee, Rio Piraparana, Schultes & Cabrera 17490 (US); 
Raudal Yayacopi, Rio Apaporis, 245 m, Schultes & Cabrera 16206 (US), 16907 (BM, US); Cachivera 
de Jirijirimo, Rio Apaporis, Schultes 12096 (BM); Schultes & Cabrera 12432 (COL, US); Soratama, 
between Rio Pacoa & Rio Kananari, 250 m, Schultes & Cabrera 12519 (US), 12778 (COL, US), 13617 
(COL, US), 1 5969 (COL, US); Garcia 13954 (BM). Between Rio Ucayali and Huallaga, Huber s.n. (P, 
type of S. brachylepis Christ). 

BRAZIL. Amazonas: type as above; Serra Curicuriari, Schultes & Lopez 9838 (BM); Serra de Sao 
Gabriel, Schultes & Lopez 8760 (BM). 

Geographical range: Amazonian Venezuela, Brazil and Colombia. 

Notes: Perhaps not really distinct from S. parkeri (No. 109), from which it is distinguished by 
having branches with indeterminate growth which root at the apex. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 309 

109. Selaginella parkeri (Hook. & Grev.) Spring in Bull. Acad. r. Belg. 10 : 146 (1843), in 

Mem. Acad. r. Sci. Lett. Belg. 24 : 226 (1850), p.p. excl. syn. Klotzsch; Baker, Fern Allies : 

104 (1887), excl. vars.; Knox in Trans, hot. Soc. Edinb., 35:254 (1950). Type from 

Guyana, Demerara: Parker s.n. (K). 
Lycopodium parkeri Hook. & Grev. in Bot. Misc. 2 : 388(1831). Type as above. 
Selaginella stellata sensu Spring in Martius, Fl. Bras. 1 (2) : 8 (1840), p.p., excl. fig. 203, non 

Fourn.(1872). 
Lycopodium geniculatum sensu J. Smith in J. Bot., Lond. 1 : 203 (1842), non Presl (1825). 
Selaginella lucidinervia Spring in Bull. Acad. r. Belg. 10 : 230 (1843). Syntypes from French 

Guiana. 
Selaginella pedata Klotzsch in Linnaea 17 : 521 (1844). Type from Guyana. 
Selaginella subarborescens Hook., Second Cent. Ferns : t.84 (1861). Type from Brazil. 
Selaginella euryclada A. Braun in Annls Sci. nat. (Bot.) V, 3 : 304 (1865). Type from Brazil. 
Selaginella euryclados A. Braun in Mber. K. preuss. Acad. Wiss. 1865 : 190, 203 (1866). 

Type as above. 
Selaginella macroclada Baker in /. Bot., Lond. 22 : 23 (1884), Fern Allies : 61 (1887). Type 

from Guyana. 
Selaginella parkeri var. pedata (Klotzsch) Baker in J. Bot., Lond. 23 : 120 (1885); Fern 

Allies : 104 (1887). Type from Guyana. 
Selaginella flabellata sensu Wright in Trans. Linn. Soc. Lond. (Bot.) II, 6 : 88 (1901), non 

Spring (1838). 

Specimens seen: 

FRENCH GUIANA. Without exact locality: Leprieur 3 (P, type of S. lucidinervia Spring); Aublet s.n. 
(BM); Karonamy, Sagot 750 (BM, NY); near R Nousineri, Leprieur 176 (K). 

SURINAM. Without exact locality: Pommier, Sara-creek, Florschiitz 280 (BM); Makami Creek, 
Lanjouw & Lindeman 3020 (BM). Coronie: Boven Maratakka, Gouggryp & Stabhel 915 (U). 
Marowijne: Nassau Mts, Lanjouw & Lindeman 2250 (BM); Marowijne R, Nassau Mts, Lanjouw & 
Lindeman 2230 (BM); Nassau Mts, Lanjouw & Lindeman 23 13 (BM). Nickerie: Timehri, Corantijn R, 
Jenman 376 (K); Corantijn R, Jenman s.n. (NY); Hostmann 880 (BM, U); Dumontier s.n. (U); 
Corantijn R, near Wonotono, Stahel & Gouggryp 2537 (U); Rombouts 72 (BM); Gran R, Hulk 253a 
(U); Nickerie R, Tulleken 482 (U, L), 362 (L), 389 (L); between Avanavero and Red Hill Falls, 
Kabalebo, Florschiitz 2104 (BM). Saramacca: Saramacca R near Yambasigado, Pulle 434 (U); near 
Pakka-pakka, Saramacca R, Florschiitz 1229 (BM); Tafelberg Creek, Saramacca R, Maguire 24104 
(BM); Tafelberg, Maguire 24813 (BM); 425 m, Maguire 24696 (BM); between Pakka-pakka and 
Ebbatop, Florschiitz 13922 (BM); Sarramacca R, Maguire 24 1 42 (BM). Suriname: Surinam R, Goddo, 
Tresling 1 92 (U), 1 79 (U); Suhosa, Suriname R, Lanjouw & Lindeman 3375 (BM). 
GUYANA. Without precise locality: type as above; Schomburgk 1 1 8 (BM, type collection of S. pedata 
Klotzch); Appun 802 (K, type of S. macroclada Baker); Mt Arisaru, Guppy 37 (BM); Macouria R, 
Jenman 2327 (BM, NY); Appun 728 (K); Rockstone, Gleason 687 (NY); Hitchcock 17320 (NY). 
Berbice: Sierra Acarai Region, 6 miles from Onoro, Guppy 244 (BM); 2nd plot \ mile from camp 1 , 
Guppy 260 (BM); 1st field by mission, Guppy 193 (BM); behind Mission, Guppy 235 (BM). Demerara: 
Anderson 244 (BM); Jenman s.n. (NY); Great Falls, Demerara R, Jenman s.n. (NY); between 
Demerara and Berbice R, De La Cruz 1593 (NY). Essequibo: near Burroburro Creek, Sandwith 1090 
(BM); Quelch & McConnell 1 1 5 (K); Kuyuwini R, A. C Smith 2552 (BM); Akarai Mts, 600-800 m, A. 
C Smith 2998 (BM); Calalebo, im Thurn s.n. (K); Pomeroon R, Jenman s.n. (NY); Marlissa crossline, 
Berbice to Rupununi Cattle Trail, Abraham 25 1 (NY); Moraballi Creek, Bartica, Richards 14 (BM, K); 
Baracara, Mazaruni, Barkly s.n. (BM); Takutu Creek, to Puruni R, Mazaruni, Fanshawe 2121 (BM); 
Tumutumari, Gleason 91 (NY), 98 (K, NY), 99 (NY), 458 (NY); Jenman s.n. (NY); Tiger Creek, 
Tumutumari, Tutin 543 (BM); Omia Creek Essequibo R, Jenman s.n. (NY); Mazaruni station, Tutin 
178 (BM); Potaro R, Jenman s.n. (NY); Junction of Mazuruni and Cuyuni R, Graham 314 (NY); 
Kurupung, Upper Mazaruni R, Leng 175 (NY), 196 (NY); along cattle-trail to Kurupukari, Beccari 
s.n.(FI). 

VENEZUELA. Amazonas: Yavita, 128 m, L Williams 14058 (BM). Bolivar: Mt Duida, 230 m, Tate 
1007 (NY); Rio Torono, above junction with Rio Paragua, 280 m, Killip 37400 (BM). Rio Paragua: 
Salto de Auraima, 275 m, Killip 37342 (BM). 
COLOMBIA. Amazonas: Rio Miritiparana, 210 m, Schultes & Cabrera 16470 (BM, US): Rio 



310 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Miritiparana, Cano Guacaya, 215 m, Schultes & Cabrera 15719 (UC). Caqueta: Florencia, Perez 
Arbelaez 630 (BM), 667 (BM); Morelia, 350 m, v. Sneidern A1263 (S). Putumayo: Umbria, 325 m, 
Klug 1711 (BM, NY), 1788 (BM, NY); Solano, 8 km SE of T res Esquinas, on Rio Caqueta below 
mouth of Rio Orteguaza, 200 m, Little 9520 (COL). Vaupes: Mitu, 250 m, Schultes & Cabrera 13962 
(COL, US), 13966 (US); Cano Teemeena, Rio Piraparana, Schultes & Cabrera 17186 (COL, US), 
17416 (COL, US); Cano Peritome, below Randal Yayacopi, Rio Apaporis, c. 230 m, Schultes & 
Cabrera 15514 (US); Soratama between Rio Pacoa and Rio Kananari, 250 m, Schultes & Cabrera 
12865 (US), 16078 (US); Randal de Jerijerimo, Schultes 12097 (BM); Cerro Isibukuri, 250-700 m, 
Schultes & Cabrera 13310 (COL, UC, US), 13329 (US), 14450A (US); Soratama, Rio Apaporis, 250 m, 
Schultes & Cabrera 12779 (COL, US), 16079 (US). 

BRAZIL. Without exact locality: Glaziou 1021 1 (C, K); Glaziou 16647 (BM). Amazonas: Manaos, Ule 
53 1 1 (K); Schwacke s.n. (BM); W bank of Manhos R, Traill 1419 (K); Manaos, Luetzelburg 220 14 (M); 
Camanaos, Rio Negros, Leutzelburg 20356 (M); Janarete, Papori, Luetzelburg 23001 (M); S Maria, 
Papori, Luetzelburg 22797 (M); 80 km N of Manaus on side trail, J.B.S. 189 (BM); Panure, Rio 
Uaupes, Spruce 2540 (BM, CGE, type of S. euryclada A. Braun); Serra Barao, Igrape da Chuva, 
Taracua, Rio Uaupes, Schultes & Pires 9056 (BM); Rio Negro, 25 km NW of Manaus, 50 m, Conant 
1161 (BM); Reserva Ducke, km 26, Manaus-Hacoatiara Road, Conant 891 (BM). Para: Rio Cumina, 
Sampaio 509 (BM); R Aripecuru, Spruce s.n. (BM, CGE). 

Geographical range: Colombia, The Guianas, Amazonian Venezuela and Brazil. 

Notes: A species characterised by having 2 or 3 strobili at the ends of its branches. This 
species sometimes develops a kind of witch's broom, whose cause is unknown. Killip 37400 
(BM) and im Thurn s.n. (K) are examples. S. macroclada Baker (Appun 802) is probably a 
prostrate, sterile branch of this species. S. pedata Klotzsch is usually smaller in leaf than S. 
parkeri but is otherwise identical. Alston maintained S. subarborescens Hook, on leaf shape 
and margin characters; also growth habit, suggesting it is broader. Somers (1978) found little 
difference except the lateral leaves lacked an upper auricle and the megaspores showed 
narrow high muri. We find the characters mixed and grade into a cline and therefore consider 
them as one taxon. It is a group that requires further study. 

110. Selaginella arthritica Alston in Archo hot. 11 : 43 (1935). Type from Costa Rica, 
Cufodontis 218(BM). 

Selaginella conduplicata Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 229 (1850), p.p. quoad 

pi. Colomb. Syntypes from Brazil, Colombia and French Guiana. 
Selaginella geniculata sensu Baker, Fern Allies : 105 (1887), p.p. quoad pi. centrali- 

americana. 
Selaginella geniculata var. conduplicata (Spring) A. Braun in Annls Sci. nat. (Bot.) V, 3 : 303 

(1865), p.p. quoad pi. Colomb. Type as above. 

Specimens seen: 

COLOMBIA. Antioquia: Villa Arteaga, Mutata, Uraba, 30-60 m, Schultes & Cabrera 18662 (US); 

Turbo, Golfo de Uraba, Schott 2 (NY); Rio Ampurrumiado, 200 m, Gutierrez & Barkley 1 7c2 1 3 (BM); 

Guapa, S of Turbo 100 m, Haught 4659 (BM); Quebrada Congo, E of Guapa, 150 m, Haught 4664 

(BM, COL). Choco: Rio Nuqui, 50 m, Haught 5460 (BM, COL). 

ECUADOR. Santiago-Zamora: near Mendez, 525-750 m, Camp E 893 (BM, NY). 

Geographical range: Colombia and south-east Ecuador, at low elevations. 

Notes: Schultes & Cabrera 1 8662 from Antioquia is a large form. 

111. Selaginella geniculata (C. Presl) Spring in Bull. Acad. r. Belg. 10 : 230 (1843); C. Presl, 
Bot. Bemerk. : 151 (1844); Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 227 (1850); A. 
Braun in Annls Sci. nat. (Bot.) V, 3:313 (1865); in Mber. K. preuss. Akad. Wiss. 
1865 : 206 (1866); Baker, Fern Allies : 105 (1887), p.p.; Alston in J. Bot., Lond. 72 : 226 
(1934); Knox in Trans, bot. Soc. Edinb. 35 : 285 (1950). Type from Peru or Ecuador (see 
Notes). 



SELAGINELLA IN TROPICAL SOUTH AMERICA 311 

Lycopodium geniculatum C. Presl, Rel. Haenk : 80 (1825); Spreng., Syst. Veg. 4 : 19 (1827). 

Type as above. 
Selaginella ferruminata Spring in Bull. Acad. r. Belg. 10 : 231 (1843), in Mem. Acad. r. Sci. 

Lett. Belg. 24 : 250 (1850). Type from Peru. 
Selaginella elongata Klotzsch in Linnaea 18 : 522 (1844). Type probably from Peru. 
Selaginella geniculata var. elongata (Klotzsch) A. Braun in Annls Sci. nat. (Bot.) V, 3 : 303 

(1865). Type as above. 
Selaginella geniculata subsp. elongata (Klotzsch) Hieron. in Engl. & Prantl, Nat. 

Pjlanzenf. 1 (4) : 7 1 2 ( 1 90 1 ). Type as above. 
Selaginella nodosa C. Presl, Bot. Bemerk. : 50 (1844); in Abh. K. Bohm. Ges. Wiss. V, 

3 : 580 (1844). Type from Peru. 
Lycopodium flabellatum sensu Kunze in Linnaea 9 : 9 (1835), non L. (1753). 

Specimens seen: 

COLOMBIA. Antioquia: Quebrada San Julian, near Argelia, 1800 m, Ewan 15780 (BM). Caldas: La 
Selva, Pueblo Rice, 1500 m, v. Sneidern 5490 (S); above Calarca, 1900 m, Alston 7778 (BM). Caqueta: 
Florencia, Perez Arbelaez 626 (BM). Cauca: W of Tambo, 1 100 m, Haught 5219 (BM); I. of Gorgona, 
Barclay 906 (BM); Teta, Lehmann BT 919 (NY). Choco: Bay ofChoco, Seemann s.n. (BM); Rio Naqui, 
300 m Haught 5495 (BM). Cundinamarca: near Bogota, 1000 m, Triana s.n. (BM). Narino: between 
Ricuarte & Diviso, Espinosa 2945 (BM); below Ricuarte, 1200 m, Alston 8449 (BM); near Altaquer, 
1000 m, Lehmann 86 (BM); Rio Puelmambi, 1540 m, Ewan 16038 (BM). Putumayo: Mocoa, 
550-730 m, Schultes & Cabrera 19046 (COL, US); between Urcusique and Mocoa, Ewan 16793 (BM); 
Rio Ticuanayoy, 1 100 m, Ewan 16805 (BM). Valle del Cauca: Quermal, Dagua Valley, Perez Arbelaez 
3206 (BM); 1300 m, Alston 7799 (BM); La Cubre, 1600-1800 m, Killip 5673 (NY). 
ECUADOR. Without precise locality, Jameson 210 (BM), 74 (NY); Andium Quitensium temperate 
forest region, Jameson 4 (NY), 60 (NY). Chimborozo: Hacienda de Licay near Huigra, Rose 22597 
(NY). Guayas: near Quillallpa, 1 50 m, Fagerlind & Wibom 682 bis (S); near Bucay, 305-393 m, Camp 
E3846 (BM); near Guayaguil, Jameson 401 (BM). Los Rios: Hacienda Clementina on Rio Pita, 
Asplund 5501 (S). Napo-Pastaza: Bimbino, banks of Rio Pacuno, lOhrs W of confluence with Rio 
Napo at Bimbino, 300 m, Whitmore 786 (BM); Archidona, Jameson 767 (BM); between Tena and 
Archidona, Asplund 9452 (S); Tena, Asplund 9420 (S); Vicinity of Ventura, Rose 23969 (NY). 
Pichincha: by Rio Toachi, near the confluence with Rio Pilaton, 915 m, Bell 218 (BM), 286 (BM); 
Santo Domingo de los Colorados, Fagerlind & Wibom 1636 (S); San Carlos des los Colorados, 
Fagerlind & Wibom 1636 (S); Hacienda Solento, near Santa Rosa, Canton Pajili. 1000 m, Mexia 6735 
(BM). Santiago-Zamora: near Mendez, 325-750 m, Camp E893 (BM); below Zamora, 900 m, Camp 
E49 (BM); between Rio Sabanilla & Cannilones Tambo, 1370-2135 m, Camp E65 (BM). 
PERU. Huanuco: Tingo Maria, Asplund 12358 (BM). Junin; San Nicolas, Pichis Trail, 1 100 m, Killip 
& Smith 26022 (BM), NY); Pangoa, Matthews 1083, (BM, type of S. ferruminata Spring). Loreto: 
between Yurimaguas and Balsapuerto, 135-150 m, Killip & Smith 28250 (BM, NY); near Tarapoto, 
Spruce 4624 (BM, CGE). San Martin: Las Cueras de los pavos, Tingo Maria, 625-1 100 m, Allard 
20532 (BM); Juan Jui, Alto Rio Huallaga, 400 m, Klug 383 1 (BM). 

Geographical range: Colombia, Ecuador and Peru to 12°S. 

Notes: Presl's type in the National Museum at Praha, is localised 'Luzon', but it must have 
been from Peru or Ecuador. Killip & Smith 14830 and 14843, both from Santander 
Colombia, have auricles on the lateral leaves, and a medium leaf size and shape similar to S. 
parkeri(No. 109). 

112. Selaginella dasyloma Alston in J. Bot., Lond. 72 : 228 (1934). Type from Colombia, 
Huila: Sombrerillo, Andre 343 (K, holotype; BM, NY, isotypes). 

Specimens seen: 

COLOMBIA. Huila: type as above. 

ECUADOR. Unrealised, in Cauca or Tolima, 600-1700 m, Alexander s.n. (BM, NY). Santiago- 
Zamora: between Hacienda Chontal and Santa Elena, 1020-1380 m, Camp E768 (BM). 

Geographical range: Colombia and Ecuador. 

Notes: This species may be distinguished from all other prostrate, articulate species by its 
large, obtuse, median leaves. 



312 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

113. Selaginella anaclasta Alston ex Crabbe & Jermy in Am. Fern J. 63 : 135 (1973). Type 
from Venezuela, Amazonas: Cerro de la Neblina, Rio Yatua, Maguire, Wurdack & 
Bunting 37063 (US; BM, fragment). 

Specimens seen: 

VENEZUELA. Amazonas: type as above. 

Geographical range: Confined to Venezuela. 

Notes: The multicellular teeth at the base of its leaves make this one of the most distinct 
species in the genus. 



114. Selaginella kunzeana A. Braun in Annl Sci. nat. (Bot). V, 3 : 296 (1865); Baker, Fern 
Allies: 62 (1887); Knox in Trans, bot. Soc. Edinb. 35:291 (1950). Type from Peru, 
Huanuco: Pampayaco, Poeppig 195 (BM, P). 

Lycopodium poeppigianum sensu Kunze in Linnaea 9:11 (1834), non Hook. & Grev. 
(1831). 

Specimens seen: 

VENEZUELA. Amazonas: Great Rapids of the Orinoco, Raudal de Atures 10 km S of Pto Ayacucho, 
1 00- 1 20 m, Maguire Wurdack & Bunting 36139 (BM). 

COLOMBIA. Antioquia: Rio Chiado, S of Turbo, 80 m, Haught 4635 (BM, COl); carretera de 
Medellin a Turbo, Cabrera 27 (COL); zona Cauchera de Villa Arteaga, Cabrera 54 (COL). Boyaca: 
Muzo Mines, 700 m, Lindig 1513 (BM). Caldas: Santa Cecilia Tatama, 800 m, v. Sneidern 5062 (S). 
Caqueta: Florencia, Perez Arbelaez 622a (BM). Cauca: 'La Gallera' Micay Valley, 1400-1500 m, 
Killip 7676 (NY, US), 7852 (NY, US). Choco: Rio Nugui, 50 m, Haught 5461 (BM, COL); La Equis, c. 
200 m, Molina & Barkley 19 Ch 085 (COL). Cundinamarca: La Palma, Triana s.n. (BM); 
1 150-1400 m, Garcia 12376 (BM), 12418 (BM); between La Palma and Pacho, 1 100 m, Haught 6092 
(BM, COL); Pandi, Herb. Nac. Colombeano 492 (US); Toraima, Lindig 1510 (P); between Cachipay 
and La Esperanza, Little 9462 (COL). Magdalena: above Onacu, 91 5 m, Santa Marta, Smith 2468 (BM, 
NY, US); Manaure, between Cordillera Central and Santa Marta, 975 m, Foster 1585 (COL). Valle del 
Cauca: valley of Rio Digua, 950 m, Alston 7805 (BM). 

ECUADOR. Chimborozo: without further location, Spruce 5675 (BM, CGE). Guayas: Salango, 
Barclay 633 (BM); Teresita, W of Bucay, 270 m, Hitchcock 20423 (US). Los Rios: near Quevedo, 
Canton Vinces, 50 m, Mexia 6618 (BM). Manabi: El Recreo, Eggers 14878 (P, S). Napo-Pastaza: 
between Banos & Mera, 4-500 m, Mexia 691 A (BM); Cashurco, Asplund 19363 (S). Santiago-Zamora: 
Rio Pante, W of Mendez, 580 m, Camp E1458 (BM). Tungurahua: between Banos and Cashurco, 
1 300—1 800 m, Hitchcock 2 1 897 (NY, US). 

PERU. Ayachucho: Estrella, between Huanta and Rio Apurimac, 500 m, Killip & Smith 22663 (BM, 
NY). Huanuco: type as above; Tingo Maria, Asplund 12140 (BM); Cueva de las Pavas, near Tingo 
Maria, 500-600 m, Ferreyra 2892 (BM); Cuchero, Dombey s.n. (P); Hacienda Mercedes, Churubamba, 
1870 m, Mexia 8197 (BM); Casapi, Matthews 1760 (BM). Loreto: Monte Campana, near Tarapoto, 
Spruce 4630 (BM, CGE, NY); between Tingo Maria and Pucallpa, 1400-1 500 m, Ferreyra 22 1 8 (BM). 
San Martin: Tingo Maria, 625-1 100 m, Allard 20361 (BM); Carpish, near Tingo Maria, Gascoyne- 
Cecil 105 (BM); Mariscal caceres, 60 km NE of Tingo Maria, 'Divisoria' pass through Cerro Azul on 
Tingo Maria-Pucallpa road, 1 500 m, R. & A. Tryon 5272 (BM). 

Geographical range: Mexico southwards to Peru. 



115. Selaginella lingulata Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 224 (1850); Baker, 

Fern Allies : 64 (1887), p.p. excel, syn. A. Braun; Knox in Trans, bot. Soc. Edinb. 35 : 285 

(1950). Type from Ecuador, Pinchincha: Jameson s.n. (BM, K). 
Selaginella intacta Baker in J. Bot., Lond. 21 : 335 (1883), Fern Allies : 60 (1887); Knox in 

Trans, bot. Soc. Edinb. 35 : 285 (1950). Type from Ecuador. 
Selaginella lindigii A. Braun in Annls Sci. nat. (Bot.) V, 3:297 (1865); Baker, Fern 

Allies: 65 (1887); Knox in Trans, bot. Soc. Edinb. 35:285 (1950). Lectotype from 

Ecuador. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 313 

Specimens seen: 

COLOMBIA. Antioquia: near Santa Elena, 2300—2500 m, Gutierrez & Barkley 17c024 (BM). Choco: 
Rio Nuqui, 50 m, Haught 5459 (COL). Cundinamarca: El Salto de Tequendama, near Bogota, 2550 m, 
Triana 696 (BM); Lindig \ 507 (BM); Ariste- Joseph s.n. (US), A 392 (US); Holton s.n. (K); Alston 7415 
(BM); Hermanos 17 (NY); 2300 m, Cuatrecasas 198 (COL); Falls of Rio Bogota, vicinity of El Salto, 
2470-2530 m, Little 7896 (COL); Salto de Tequendama, 2100 m, Alston 7415 (BM); Barroblanco, 
Andre 1483 (K), 1388 (K). Putumayo: between Lake Cocha and Santiago, 2350 m, Alston 8344 (BM); 
near Lake Cocha, 2700 m, Alston 8321 (BM). Tolima: Quindio Road, 3000 m, Alston 1111 (BM); 
Quebrada Peralis, Quindiva Road, 2600 m, Alston 7751 (BM). Vallee del Cauca: Cordoba, Dagua 
Valley, 80-100 m, Killig 5047 (US). 

ECUADOR. Without precise locality: Spruce 4798 (BM, CGE, NY; lectotype of S. lindigii A.Br.); 
between Rio Gamolotal and Rio Norcay, 1095-1370 m, Steyermark 52854 (BM); Andes Quitense, 
Spruce 5603 (CGE, NY). Carchi: between El Pun and Puenta Chingual, Asplund 16982 (S). Guyas: 
Guayaquil, Jameson s.n. (BM); San Nicolo, Sodiro 71 (K, type of S. intacta Baker). Napo-Pastaza: 
Baeza, 2000—2200 m, Balslev & Madsen 10318 (BM). Pichincha : type as above; Salvador, 2440 m, 
Bell 667 (BM); Aloag, El Corazon, 2865 m, Wiggins 10694 (BM); Nono Gualea, Sodiro s.n. (NY); 
woods on the western side of Pichincha, Jameson 43 (K); Corazon, Sodiro s.n. (K). Santiago-Zamora: 
between Campanas and Arenillas, along Rio Tinta, 10 leagues SE of El Pan, 2195 m, Steyermark 
53617 (BM). 

PERU. Without exact locality: Pichis Highway, Bradley 66666 (US). Huanuco: near Puente Durand, 
between Huanuco and Tingo Maria, 1500-1 600m, Ferreyra 2937 (BM). Junin: Chanchamayo Valley, 
Schunke 788 (US); Schunke Hacienda, above San Ramon, 1800m, Schunke A249 (US); 1300-1 700m, 
Schunke A25 1 (US); Pichis Trail, San Nicolas, c. 1 100m, Killip & Smith 26056 (BM). 
BOLIVIA. Without exact locality: Songo, Bang 909 (US). La Paz: Ticunhuaya, 1525 m, Tate 1076 
(US). 

Geographical range: Colombia south to Bolivia. 

Notes: Somers (1978) considered the type of this species to be atypical, having nearly 
elliptical and widely spaced lateral leaves which could be indicative of shade growth, which 
grades into the form hitherto called S. lindigii A. Braun. Similarly Somers (l.c) regarded the 
type of Baker's S. intacta to be an extremely depauperate form of S. lingulata, often confused 
with S. atirrensis (No. 120), but lacking the multicellular cilia of that taxon. Archer 1517, 
from Colombia, is typical, having the auricles of the median leaf somewhat unequal, and 
with scant cilia on the axillary leaf bases. 

116. Selaginella articulata (Kunze) Spring in Flora, Jena 21 : 182 (1838), in Annls Sci. nat. 
(Bot.) II, 11 : 228 (1839), in Bull Acad. r. Belg. 10 : 229 (1843), Mem. Acad. r. Sci. Lett. 
Belg. 24 : 21 1 (1850); Baker, Fern Allies : 82 (1887); Knox in Trans, bot. Soc. Edinb. 35 : 
283 (1950). Type from Peru, Loreto: Maynas Alto, Poeppig s.n. (K, P). 

Lycopodium articulatum Kunze in Linnaea 9:10 (1835), Farrnkr. 1 : 243 (1847). Type as 
above. 

Specimens seen: 

COLOMBIA. Putumayo: above Mocoa, 600 m, Ewan 16710 (BM). 

ECUADOR. Napo-Pastaza: Archedonia, Jameson 714 (BM); Mera, Asplund 18781 (S); near Tena, 

400 m, Mexia 7219 (BM); between Puyo and Canelos, 325-375 m, Mexia 6842 (BM); near Puyo, 

Fagerlind & Wibom 1 185 (S); Mera, 1200 m, Ollgaard & Balslev 9096 (AAU, BM); near Hacienda 

Ortiz, 18 km from Puyo on road to Tena, 1 100 m, Ollgaard & Balslev 9264 (AAU, BM). 

PERU. Loreto: type as above; Monte Campana, near Tarapoto, Spruce 4627 (BM, CGE, NY). 

Geographical range: Colombia, Ecuador and Peru. 

Notes: Distinguished from the other two pubescent articulate species (Nos. 104 and 105) in 
having rather close, spreading, obtuse leaves on the main stem. S. sericea (No. 128) is a very 
similar species, but the stems are always glabrous and the lateral leaves have two auricles. 

117. Selaginella valida Alston in J. Bot., Lond. 70:281 (1932). Type from Brazil, Sao 
Paulo: S Jose do Barreiro, Hoehne & Gehrt 1 7698 (BM). 



314 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Specimens seen: 

BRAZIL. Sao Paulo: Santos, Mosen 3812 (P); type as above. 

Geographical range: So far recorded only in Sao Paulo. 

118. Selaginella calceolata Jermy & Rankin sp. nov. (Figs 1 7 & 18) 

Selaginella arenaria Baker in J. Bot., Lond. 21 : 82 (1883), p.p. non spec, in herb. Kew. 
Planta caulibus prostratis ramis ascendentibus, foliis intermediis calceolatis auricula 
exteriore magna distincta. 

Caules prostrati articulati pallidi straminei; rhizophoris dorsaliter exorientibus; ramis 
paucis, primariis 3-6 cm longis, alternis, saepe suberectis, ultimis 1-2 cm longis, patentibus, 
Folia ubique heteromorpha; folia lateralia 1-8-2-2 mm longa, 0*8-1-1 mm lata, oblongo- 
elliptica vel oblongo-lanceolata, apice obtuso vel subacuto, semifacie superiore denticulata, 
basi rotundata, semifacie inferiore basi truncata apicem versus denticulata, aliter integra; 
folia axillaria 1-8-2*0 mm longa, c. 0*7 mm lata, oblongo-elliptica, denticulata, apice 
rotundato vel late obtuso, basi plus minusve cuneata; folia intermedia 1-5-1-8 mm longa, 
0- 5-06 mm lata, adpressa, imbricata, calceolata, auricula magna exteriore calceum 
faciente, apice acuto vel subacuto, integri, margine inconspicuo. Strobili 1-2 mm longi, 
basi feminei, supra masculi; megasporophylla c. 1 *5 mm longa, 0*7 mm lata, late ovata, apice 
denticulate acuminato vel acuto; megasporae reticulatae; microsporophylla 0*8 mm longa, 
0*4 mm lata, megasporophyllo similia sed manifestius carinata; microsporae non visae. 

Typus: Colombia: Puerto Lopez, Intendencia del Meta, in ripa humida umbrosa, 240 m, 27 
July 1944, E.L. & R.R. Little 8241 (COL, holotype; BM, US, isotypes). 

Other specimens seen: 

COLOMBIA. Boyaca: Orucue, Los Llanos, Rio Meta, 140 m, Cuatrecasas & Barriga 4389 (COL). 

BRAZIL. Amazonas: on Rio Vaupes at Panure, Spruce 286 1 (BM). 

Geographical range: Eastern Colombia to western Brazil. 

Notes: Spruce's gathering was mixed, and only the specimen at Kew agrees with Baker's 
description of S. arenaria (No. 78), thus necessitating the description of this species. We are 
grateful to Paul Somers for drawing attention to this situation. 

119. Selaginella marginata (Humb. & Bonpl.) Spring in Flora, Jena, 21 : 194 (1838), in 
Martius, Fl. Bras. 1 (2) : 1 17 (1840), ex Decne in Arch. Mus. 2 : 192 (1841-42), in Bull. 
Acad. r. Belg. 10 : 229 (1843), in Mem. Acad. r. Sci. Lett. Belg. 24 : 21 1 (1850), p.p. excl. 
loc. mexicanis; A. Braun in Annls Sci. nat. (Bot.) V, 3 .291 (1865); Baker, Fern Allies : 61 
(1887). Type from Venezuela, Apure: mouth of R Meta, Humboldt s.n. (B). 

Lycopodium marginatum Humb. & Bonpl. ex Willd., Spec. PI. 5 : 41 (1810); Kunth, Nov. 

Gen. Spec, 1 : 391 (1816), Syn. PI. 1 : 96 (1822); Sprengel, Syst. Veg. 4:18 (1827). Type as 

above. 
Selaginella excurrens Spring in Martius, Fl. Bras. 1 (2) : 128 (1 840). Type from Brazil. 
Selaginella marginata subsp. distorta Spring in Flora, Jena 21 : 196 (1838). Type from 

Brazil. 
Selaginella marginata var. minor Spring I.e. Type from Brazil. 
Selaginella distorta (Spring) Spring in Bull. Acad. r. Belg. 10 : 229 (1843). 
Selaginella distorta var. minor (Spring) Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 2 1 3 

(1850). 
Selaginella distorta var. major Baker in J. Bot., Lond. 21 : 335 (1883). Type from Brazil. 
Selaginella chromatophylla Silveira in Bolm Comm. geogr. geol. Minas Geraes 5 : 124 

(1898) and vars. Types from Brazil. 
Selaginella urbanii Hieron. in Engl. & Prantl, Nat. Pjlanzenf 1 (4) : 709 (1901). Type from 

Brazil. 
Selaginella moseni Hieron. I.e. : 709. Type from Brazil. 
Selaginella burchellii Hieron. I.e. : 709. Type from Brazil. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 



315 



** 



9 






i 










4wS 



1* 



1 




& 



X*^- 



/ ; 






Det. AC. Jenny & J.M. Rankin 



fela 



TYPE SPECIMEN 

or 
A 



«fi"««<\ c^leeeledc 



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PLANTS OF COLOMBIA, SOUTH AMERICA 

Ban 

''elt^lnella »p • 

(Cannot be named «t jre««rit.) 
* f ;retifcdorrs* 

I'olat da^>, ahaced soil und*r thick ft. 



- Veto 
27, ^WftC 

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Maxon & Morton, IS* 6. 

Fig. 17 Selaginella calceolata Jermy & Rankin: Type specimen, EL. & R.R. Little 824 1 (COL). 



316 



A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 




Fig. 18 Selaginella calceolata Jermy & Rankin: A. Close-up of lateral (/) and axillary (a) leaves, 
x 25. B. Close-up of median leaf, x 56. Both from E.L. & R.R. Little 8241. 

Selaginella excurrens var. glazioviana Hieron. ex Rosenstock in Hedwigia 46 : 166 (1907). 

Type from Brazil. 
Selaginella breuensis Silveira in Fl. Serras Mineiras : 79 (1908). Type from Brazil. 
Selaginella herteri Hieron. in Reprium Spec. nov. Regni veg. 13 : 421 (1914). Type from 

Uruguay. 
Selaginella caracensis Goebel in Flora, Jena 108 : 3 1 6 (19 1 5). Type from Brazil. 
Selaginella ypirangensis Hieron. ex Herter in Beih. hot. Zbl. 39 : 253 (1922). Type from 

Brazil. 
Selaginella ypirangensis var. riograndensis Herter I.e. : 256. Type from Brazil. 

Selaginella wielewskii Hieron. ex Rosenstock in Hedwigia 43:234 (1904). Type from 
Brazil. 



Specimens seen: 

VENEZUELA. Without exact locality: Grosse s.n. (BM). Apure: type as above. Aragua: San Matos, 
Famayo 633 (BM). Guarico: between San Juan de los Morros and Parapara, 400 m, Alston 5997 (BM), 
6004 (BM). 

BOLIVIA. Santa Cruz: S Cruz, 422 m, Tutin 1 549 (BM); 460 m, Brooke 5844 (BM). 
BRAZIL. Without exact locality: Glaziou 15802 (C, P), 22646 (C), 7355 (BM, C), 7969 (C), 15803 
(C), 12296 (BM, C); Langsdorf s.n. (K); Sellow s.n. (K); Burchell 8724 (K); Dusen 3726 (BM). Bahia: 
Blanchet s.n. (M). Goias: 25 km N of Brasilia, 950 m, Irwin, Souza & Reis dos Santos 1 1282 (BM). 
Maranhao. Una dos Botes, Carolina Rio Tocantins, Pires & Black 2037 (BM). Mato Grosso: Corumba, 
Robert 728 (BM); Hoehne 45 (BM); Rasteira s.n. (BM); Sierra de Chapada, Moore 85 (BM). Minas 
Gerais: without further locality; Glaziou 201 16 (C); Claussen s.n. (P), St Hilaire 272 (P); Conselheira 
Matta-Rodeador, Brade 13961 (BM); Campanha, Schreiner s.n. (BM); Rio Grande do Sus, Caldas, 
Mosen s.n. (BM); Caldas, Mosen 20 1 8 (P), 20 1 9 (C, P), 2020 (P), 202 1 (C, P), 2023 (C); Santa Barbara do 
Matto Dentro, Hoehne 5004 (BM); Casa de Pedra, Tiradentes, Silveira 160 (BM); Serra do Cipo, 
Silveira 395 (BM); Serra do Papagaio, Silveira 152 (BM); Palmeiras, Lindman A 2499 (NY); Cuyaba, 
Lindman A 2347 (K). Parana: Schwacke s.n. (BM); between Cuntiba and Butiatuba, Schwacke Coll. II 
No. 37 (BM), 2706 (BM, P); Serra do Mar, Ypiranga, Dusen 3726 (BM, P); Lucena, Wielewski 7 (NY); 



SELAGINELLA IN TROPICAL SOUTH AMERICA 317 

Rio Negro, Annies 93 (NY); Serrimba, Dusen 15188 (BM); Serra do Mar, Banhado, Dusen 1 4465 (BM); 
Curitiba, Augley & Mattos 3831 (UC). Piaui: Netto 80 (BM). Rio de Janeiro: without further locality, 
Raben s.n. (BM, C), Santa Magdalena Pedra das Flores, Lima & Brade 13190 (BM); Serra dos Orgaos, 
1000 m, Brade 1681 1 (BM); Sto Antonio, Glaziou 7355 (BM). Rio Grande du Sol: Villa Germania, 
Jurgens L 23 (BM, NY); Jurgens & Stier 310 (BM); Cachoeira, Lindman A 1153 (BM); Nova 
Wuertemberg, Jurgens & Stier 3 1 1 (BM); Taquara, Caracol, Dutra 1612 (BM); S Antonio da Patrulha, 
Dutra 134 (BM); Bomjesus, Dutra 235 (BM), 1608 (BM); S Francisco de Paula, Rambo 44799 (BM), 
46312 (BM), 45638 (BM); Aparados da Rocinha near Bom Jesus, Rambo 45363 (BM); Toca do Tigre 
near Itapoan, Rambo 48844 (BM). Santa Catarina: Lages, Rambo 49548 (BM); Spannagel 103 (NY), 
235 (NY), 417 (BM); Vaccao Gordao, Lages, Spannagel 45 (NY). Sao Paulo: Campos do Jordao, Porto 
3114 (BM); 1600 m, Leite 3658 (BM); Lorena, Luederwaldt s.n. (BM). 

Geographical range: Mexico, southwards to Uruguay and northern Argentina. 

120. Selaginella atirrensis Hieron. in Engl. & Prantl, Nat. Pflanzenf. 1 (4): 711 (1901); 
Knox in Trans, hot. Soc. Edinb. 35 : 290 (1950). Type from Costa Rica, Cartago: near 
Atirro, 650 m, Bonn Smith 5103 (NY, US). 

Selaginella intacta sensu auct., non Baker (1883). 

Specimens seen: 

COLOMBIA. Cauca: Cordillera occidental, 1600 m, Bischler 1497 (COL). Choco: bank of Rio San 

Juan, near Andagoya, c. 60 m, Killip 35379 (COL); between La Oveja and Quibdo, Archer 1666 (BM). 

Narino: Lehmann 88 (BM). Valle del Cauca: Rio Calima, between Pailon and El Coco, 50 m, 

Cuatrecasas 21244 (BM); km 99 on Cali-Buenaventura Highway, c. 200 m, Haught 5319 (BM, COL); 

Cordoba, Dagua Valley, Killip 5048 (US), 5032 (US). 

PERU. Junin: Chanchamayo valley, 1500 m, Schunke 229 (BM); Schunke Hacienda, above San 

Ramon, 1300-1700 m, Schunke A2 51 (BM); 1 800 m, Schunke A249 (BM). 

Geographical range: Costa Rica south to Colombia. 

121. Selaginella kraussiana (Kunze) A. Braun, Appl Ind. Sem. Hort. Berol : 22 (1860); 
Baker, Fern Allies: 65 (1887). Type from South Africa: Zitzikamma District, Kraus 
Marks s.n. (? B). 

Lycopodium kraussianum Kunze in Linnaea 18 : 1 14 (1844). Type as above. 

Selaginella hortensis Mett., Fil. Hort. Lips. : 125 (1856). Type ex hort. Lips. 

Selaginella canescens Fee, Crypt. Bres. 2 : 99 (1873); Alston in Reprium Spec. nov. Regni 

Veg. 40 : 318 (1936). Type from Brazil. 
Selaginella denticulata Hort.; Jl R. hort. Soc. 8 : 35, 123 (1887), non (L.) Link (1841). Type 

from Canarias. 

Specimens seen: 

BRAZIL. Rio de Janeiro: Glaziou 4489 (C, K; type of S. canescens Fee), 12297 (C, K). Rio Grande do 
Sul: S Cruz, Jurgens L 35 (BM). Santa Catarina: Joinville, Schmalz 149 (BM, NY); Cantareira, Usteri 
s.n. (BM). 

Geographical range: Native to Africa and Macaronesia but escaped from cultivation or 
accidentally introduced in the New World from Virginia to Chile. 

Notes: This was described by Fee as an endemic species, S. canescens, which, as far as we can 
see, is conspecific with S. kraussiana. The general assumption is that it has been introduced 
in South America, but it is conceivable that it is native. It is distinct from other articulate 
species, all of which are confined to the neotropics. 

122. Selaginella asplundii Alston ex Crabbe & Jermy in Fern Gaz. 11 : 257 (1976). Types 
from Peru, Huanuco: Carpish, 2700 m, Asplund 12822 (S, holotype; BM, isotype). 

Specimens seen: 

COLOMBIA. Cauca: Valle de Las Papas, near Valencia, 29 1 m, Idrobo, Pinto & Bischler 3643 (BM). 

PERU. Huanuco: type as above; ibid. loc. 2650 m, Asplund 12843 (S). 

Geographical range: Colombia and Peru. 



318 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

123. Selaginella epirrhizos Spring in Bull. Acad. r. Belg. 10 : 229 (1843), in Mem. Acad. r. 
Sci. Lett. Belg. 24 : 218 (1850); Baker, Fern Allies : 81 (1887); Alston in Pulle, Fl. Surin. 1 : 
167 (1938); Knox in Trans, bot. Soc. Edinb. 35 : 284 (1950). Type from French Guiana, 
Cayenne: Leprieur 12 (P). 

Selaginella rigida Linden, Catalogue 8:15 (1853), nom. nud.; A. Braun, Index Sem. Hort. 

bot., Berl. Appendix 1857 : 22 (1857). 
Selaginella affinis A. Braun in Annls Sci. nat. (Bot.) V, 3 : 296 (1865); Baker, Fern Allies : 63 

(1887). Type from French Guiana. 
Selaginella poeppigiana var. guyanensis Spring in Mem. Acad. r. Sci. Lett. Belg. 24 : 2 1 8 

(1850). Type from French Guiana. 
Lye op odium plumosum sensu Aublet, Hist. PL Guian. : 967 (1775), non L. (1753). 

Specimens seen: 

FRENCH GUIANA. Without precise locality: Aublet s.n. (BM); Leprieur s.n. (K, P); type as above. 
Guyane: Bradel cataract, Rio La Comte, Leprieur 178 (P); Serra de Ouanari, Black, Vincent & DAage 
1 7638 (BM, IAN). 

SURINAM. Without precise locality: Hostmann & Kappler 3 (L). Nickerie: Upper Nickerie R, 
Tulleken 509 (L); Phaedra, Voltze s.n. (U). Sarramacca: Bergendaal, Focke 386 (U); between Pakka- 
pakka and Ebatop, Florschutz 1393 (BM). Suriname: Blaauwe Berg, Splitgerber 893 (L). 
GUYANA. Without exact locality: Penal Settlement, Guppy 2 (BM); between Demerara and Berbice 
Rivers, De La Cruz 1606 (NY). Demerara: Malali, Demerara R, De La Cruz 2679 (K, NY); Demerara, 
Andersen 40 (BM). Essequibo: Kamuni Creek, Groete Creek, Essequibo R, Maguire & Fanshawe 
22847 (BM); Smith 2108 (BM); Mt Arisaru, Guppy 28 (BM); Kaieteur Falls, Appun s.n. (BM); Head 
Falls, Mazaruni R, Jenman 645 (K), s.n. (NY); Mazaruni Station, Sandwith 1231 (BM); Kamakusa, 
Upper Mazaruni R, De La Cruz 2793 (NY); Camaria road, junction of Mazaruni and Caryuni Rivers, 
Graham 368 (BM, NY), 407 (BM, NY); near Lower Camaria Tutin 147 (BM); Richards 841 (BM, K); 
Bartica, Hitchcock 17257 (NY); Bartica Grove, Jenman 2326 (BM); Arawak Matope, Cuyuni R, Tutin 
382 (BM); Rockstone, Gleason 685 (NY); Tiger Creek, Potaro R, Jenman s.n. (NY); Potaro Road, 
Guppy 7 (BM); Tumutumari, Gleason 57 (NY), 455 (NY), 456 (NY); Hitchcock 17379 (NY); Linden 29 
(NY); Tutin 547 (BM); Kurupung, Mazaruni R, Leng 270 (NY); Sierra Acarai region, c. 3 km NE of 
Camp 1 , Guppy 335 (BM); Takutu Creek to Puruni R, Mazaruni R, Fanshawe 1 1 89 (NY), 2033 (BM); 
Kamuni Creek, Groete Creek, Essequibo R, Fanshawe 1821 (NY); Baracara, Mazuruni R, Barkly s.n. 
(BM); Upper Mazaruni, De la Cruz 2333 (NY), 2236 (NY), 2401 (NY); near Dadanawa, Upper 
Rupununi, De La Cruz 1811 (NY); Huradiaiah, Moruka R, Pomeroon District, De La Cruz 1019 
(NY); Pomeroon R, Jenman s.n. (NY). 
BRAZIL. Amazonas: S Francisco, Rio Acre, Ule9\4\ (K). 

Geographical range: The Guianas, extending into NW Brazil. 

Notes: A form with smaller leaves occurs in Guyana. It has been seen from Tiger Creek, 
Potaro R, Jenman s.n. (NY); Tumatumari, Potaro R, Hitchcock 17379 (NY); Tiger Creek, 
Tumatumari, Tutin 544a (BM). The species has a characteristic appearance due to the 
imbricate leaves of the tapering terminal branchlets and regularly spaced lateral leaves of the 
main stem. 

124. Selaginella poeppigiana (Spring) Spring ex Splitg. in Tijdschr. Natuurl. Gesch. Physiol. 
7 : 443 (1840), in Mem. Acad. r. Sci. Lett. Belg. 24 : 217 (1850), p.p. quoad pi. andina; 
Baker, Fern Allies : 62 (1887), p.p.; Knox in Trans, bot. Soc. Edinb. 35 : 286 (1950). Type 
from Ecuador, Pichincha: Jameson s.n. (K). 

Selaginella sulcata subsp. poeppigiana Spring in Flora, Jena 21:185 (1838). Type as above. 
Lycopodium poeppigianum Hook. & Grev. in Bot. Misc. 3 : 106 (1833), p.p., non I.e. 2 : 
393, No. 144(1831). 

Specimens seen: 

ECUADOR. Pichincha: type as above. Santiago-Zamora: Quebrada Honda, Andre K 456 (K, NY); 

between Quebrada Honda and Tambo Valladolid, 2000-3000 m, Steyermark 54592 (BM). 

PERU. Ancash: Pampayacu, Kanehira 182 (US). Cuzco: Paucartambo, Marcachea, near Achirani, 

Vargas 11147 (UC); Paucartambo, between Patria and Libertad, 650 m, West 7102 (UC). Junin: 



SELAGINELLA IN TROPICAL SOUTH AMERICA 319 

Schunke Hacienda, above San Ramon, 14-1700 m, Schunke A 252 (BM, US), A 253 (BM); Killip & 
Smith 24657 (BM, NY); Chanchamgyo Valley, Schunke 192 (BM), 231 (BM); Pangoa, Matthews 1085 
(K). Loreto: Monte Campana, near Tarapoto, Spruce 4626 (BM, CGE, NY). 

Geographical range: Ecuador to Peru. 

Notes: This species, because of the auricles on the lateral leaves, may perhaps be regarded as 
conspecific with S. sulcata (No. 129). If maintained as distinct this epithet presents a 
complicated problem of nomenclature. Hooker & Greville originally applied the name L. 
poeppigianum to a Cuban plant collected by Poeppig, but in 1833 included this species 
under L. stoloniferum. They write k We find, upon examining authentic specimens of the L. 
stoloniferum of Swartz, in the Banksian Herbarium [i.e. in the British Museum], communi- 
cated by himself from Hispaniola, that we have committed an error in regard to the present 
plant. The species we named L. Pceppigianum is the true plant of Swartz, and was indeed 
transmitted to us as such by our friend Professor Kunze. The specific appellations of the two 
plants will, therefore, have to be transposed, and the synonyms of our L. stoloniferum 
referred to what we have erroneously called L. Pceppigianum". 

Hooker had two specimens from 'Demerara, Parker & Pichincha, Jameson". In 1834 
Kunze (Linnaea, 9:11) referred a plant from Pampayaco, Poeppig to this species; this has 
since been taken out as S. kunzeana (No. 114). In Flora, Jena 21 Spring makes it a 
subspecies of S. sulcata and notes three specimens: Peru: Poeppig, Jameson and Demerara: 
Parker. In his monograph, Spring made two varieties: mexicana (= S. galeottii Spring) and 
guyanensis (= S. epirrhizos, No. 123) evidently regarding the Andine plant as the type, which 
reduces the type to Jameson' 's specimen in Hooker's herbarium (i.e., K). 

125. Selaginella suavis (Spring) Spring in Bull. Acad. r. Belg. 10 : 229 (1843), Mem. Acad. r. 
Sci. Lett. Belg. 24 : 216 (1850). Type from W. Brazil, without exact locality, Sellow s.n. (B, 
K). 

Selaginella sulcata subsp. suavis Spring in Flora, Jena 21 : 185 (1838); in Martius & Endl., 

Fl. Bras. 1 (2) : 1 3 1 ( 1 840). Type as above. 
Selaginella glaziovi Fee, Crypt. Bres. : 232 (1869); I.e. Suppl. : 101 (1873). Type from Brazil. 

Specimens seen: 

PERU. Junin: Tarma, San Ramon, Valle de Chanchamayo, 800-900 m, Constance & Tovar 2269 

(UC). 

BRAZIL. Without exact locality: type as above (BM fragment = S. sulcata (No. 129) [det. Somers]). 

Espiritu Santo: Serra da Caparao, 1550 m, Mexia 4072 (BM). Mato Grosso: Esperidras, Hoehne 733 

(BM). Minas Gerais: Cel-Pacheco, Heringer 1522 (BM). Rio de Janeiro: Glaziou 4482 (C, K), 4502 

(C, K); Corcovado, Didrichsen 3135 (C); Monte Serrat, near Itatiaya, Rose & Russell 20440 (NY, P); 

Itatiaia, Dusen 654 (BM); Brade 10048 (BM); Santa Magdalena, 1000 m, Lima & Brade 13189 

(BM). Santa Catarina: Schreiner s.n. (BM). Sao Paulo: Serra da Cantareira, Loefgren 530 (BM), 12 (C, 

S), 168(P). 

Geographical range: Brazil and reaching central Peru in the west. 

Notes: There is confusion over the typification of this taxon. Spring (1838 : 185) refers to 
Lycopodium marginatum of Gaudichaud in Freycinet, Voyage Autour Monde, Bot. : 286 
(1828) which refers to a Brazilian plant (Rio de Janeiro: calcada da Estrella). Gaudichaud's 
specimens at K (fide Alston (1936) et nostra) and at B (Somers, pers. comm.) is S. sulcata 
(No. 129). In Fl. Bras. Spring began to emend the description, mentioning L. marginatum 
[sensu] Gaud, in synonymy, but lists other Brazilian specimens the first of which is Sellow (in 
Brazilia meridionali) and may be selected as type, although the fragment at BM suggests a 
mixed gathering and further adds to the confusion. 

126. Selaginella silvestris Asplund in Ark. Bot. 20 A : 30 (1926). Type from Bolivia, La Paz: 
El Chaco, Sur Yungas, Asplund 1 1 40 (BM (fragment); UPS; Herb Birger). 

Selaginella poeppigiana sensu Baker, Fern Allies : 62 ( 1 887), p.p. non Spring ( 1 838 & 1 840); 
Hieron.tfo/. 76.34:581 (1905). 



320 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

Specimens seen: 

COLOMBIA. Antioquia: Palmitas, Skolnik, Daniel & Barkley 19 An 158 (BM). Caldas: Rio Santa 
Rita, Salento, 1600-1800 m, Killip & Hazen 8991 (NY), 10137 (NY). Caqueta: Balsillas Mts, Ariste- 
Joseph s.n. (US). Cauca: Canaan, Mt Pirace, 3100-3300 m, Pennell & Killip 6687 (BM, NY); near 
Popayan, 1800 m, Alston 8023 (BM). Cundinamarca: Tocarema, 2500 m, Lindig 1508 (BM); between 
La Vega and Facatativa, 2100 m, Haught 6163 (BM); La Esperanza, Perez Arbeldez 2576 (COL). Norte 
de Santander: Paramo de Fontibon, Pamplona, 2400 m, Alston 7128 (BM); La Mesita, Pamplona, 
2725 m, Alston 7235 (BM); Loso, N of Toledo, 2200-2400 m, Killip & Smith 20402 (NY). Putumayo: 
near Lake Cocha, 2700 m, Alston 8309 (BM); near Llorente, E of Quebrada Balsal, 2290 m, Ewan 
16260 (BM); Valle de Sibundoy, 1-5 km NE Sibundoy, c. 2200 m, Bristol 443 (COL). Santander: near 
La Baja, 3000 m, Killip & Smith 18114 (NY), 18357 (NY). Tolima: Quebrada Peralis, Quindin Road, 
2600 m, Alston 7753 (BM). 

ECUADOR. Without precise locality: Spruce 4785 (BM, CGE, NY). Azuay: near Servilla de Oro, 
2440-2745 m, Camp E 4285 (BM). Guyas: 3 km W of Bucay, Teresita, 270 m, Hitchcock 20423 (NY). 
Manabi: Monte Christi Hill, Fagerlind & Wibom 509 (S). Napo-Pastaza: near La Chonta, Rose 23467 
(NY); road Baeza-Lago Agrio, 23 km from Baeza at Rio Oyacachi, 1500 m, Balslev & Madsen 10451 
(BM). Pichincha: Nono-Gualea, Sodiro s.n. (NY); Tandapi, 2000 m, Mille s.n. (NY); Quito, Jameson 
28 (BM); Cerro Carazon, Camp E 1667 (BM); San Florencio, 1580 m, Andre 3655 (NY); Pichincha, 
Sodiro s.n. (NY); Chaupi-Sagcha, Pululagua, 1830 m, Bell 436 (BM). Tungurahua: Cusatanga near 
Ambato, Pachano 1 76 (US). 

PERU. Cuzco: Keros Valle Cosnipata, 950 m, Scolnik 895 (BM); between Mistiana and Keros, 650 m, 
Vargas 7398 (BM); Valle de Santa Anna, prov. Convencion, Herrera 3009 (US); Convencion, Potrero, 
8m W of Quillabamba, 1200m, R. & A. Tryon 5394 (BM). Huanuco: Carpish Divide, 2650m, 
Sandeman 51 18 (BM); Asplund 12842 (S). Junin: Perene River, 1 190 m, Gascoyne-Cecil 90 (BM). 
BOLIVIA. La Paz: type as above; Sirupaya near Yanacadis, Sur Yungas, 1650 m, Buchtien 441 (US); 
San Felipe Unduavi, 2650 m, Buchtien 1 1 367 (US). 

Geographical range: Guatemala southwards to Bolivia. 

Notes: This species is very close to S. trisculcata (No. 127), but the oblong lateral leaves are 
rounded at the apex, and only toothed towards the base. 

127. Selaginella trisulcata Asplund in Ark. Bot. 20A : (1926); Knox in Trans, hot. Soc. 

Edinb. 35:286 (1950). Type from Bolivia, La Paz: El Chaco, Sur Yungas, 1900 m, 

Asplund 1482 (S, holotype; UPS, Herb Birger, isotypes). 
Selaginella buchtienii Hieron. in Meded. Rijks-Herb 27 : 2 (19 1 5), nom. nud. 
Selaginella poeppigiana war. peruviana A. Braun in Annls Sci. nat. (Bot.) V, 3 : 295 (1865). 

Specimens seen: 

ECUADOR. Napo: Cuyuja, S of Rio Papallacta, 2500-2650 m, Balslev & Madsen 10540 (AAU, BM). 
PERU. Ayacucho: Aina, between Huanta and Rio Apurimac, 750-1000 m, Killip & Smith 2275 1 (BM, 
NY); Carrapa, between Huanta and Rio Apurimac, 1500 m, Killip & Smith 22359 (BM, NY). Cuzco: 
San Miguel, Urubamba Valley, 1800—2300 m, Herrera 2024 (BM); Cook & Gilbert 878 (NY, US); 
Aobamba, Huadquina, prov. Convencion, Vargas 8154 (BM). Huanuco: Carpish Pass, 2750 m, Allard 
20996 (BM). Junin: Oxapampa, prov. Pasco, 1600 m, Soukup 3359 (BM); Carpapata, above 
Huacapistana, 2400 m, Killip & Smith 24373 (BM, NY); 2700-3200 m, Killip & Smith 24467 (BM, 
NY); Huacapistana, 1800 m, Killip & Smith 24309 (BM, NY); Schunke Hacienda, above San Ramon, 
1300-170 m, Schunke H 253 (US); Chanchamayo, Schunke s.n. (BM); Pichis trail, Yapas, 
1350-1600 m, Killip & Smith 25588 (BM, NY). 

BOLIVIA. Without precise locality: Bridges s.n. (BM); Bang 51 (BM), 66 (BM). Cochabamba 
Antahuanca, Quebrada del Rio Espiritu Santo, 750 m, Buchtien 2255 (US); ed. Rosenstock 92 (BM) 
Incahaca, 2440 m, Brooke 6659 (BM). La Paz: type as above; near Yungas, Bang 577 (BM, NY, US) 
1830 m, Rusby 452 (NY, US); Polo-Polo, near Coroico, N Yungas, 1000 m, Buchtien 3550 (US); ed. 
Rosenstock 101 (BM), 4847 (US); Unduavi, 2440 m, Rusby 454 (NY); Yungas, Caranavi, Ellenberg 
6223 (AAU); Sur Yungas, Chulumani, 2135 m, Brooke 653 1 (BM). 

BRAZIL. Para: vicinity of Cachoeira, km 96, road BR22, Capanema to Maranhao, Varzea forest, 
Prance & Pennington 1 866 (BM). 

Geographical range: Ecuador, Peru, Bolivia, with an outlier in mid-Brazil. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 321 

128. Selaginella sericae A. Braun in Annls Sci. nat. (Bot.) V, 3 : 298 (1865); in Mber. K. 
preuss. Akad. Wiss. 1865 : 187 (1866); Baker, Fern Allies : 82 (1887). Type from Ecuador, 
without exact locality: Andes of Quito, Cuming 82 (B). 

Selaginella sericea var. latifolia A. Braun in Annls Sci. nat. (Bot.) V, 3 : 298 (1865). Type 

from Ecuador. 
Selaginella sericea var. elongata Sodiro, Rec. Crypt. Vase. Prov. Quito : 94 (1883), Crypt. 

Vase. Quito : 613 (1893). Type from Ecuador. 

Specimens seen: 

ECUADOR. Without precise locality: type as above; Spruce 4786 (BM, CGE, NY); Jameson s.n. (K); 
Fraser s.n. (BM). Guayas: between Hacienda Chontal and Santa Elena, 1035-1400 m, Camp E812 
(BM). Pichincha: Nanegal, Sodiro s.n. (K); Mondo, Sodiro s.n. (K); San Florencio, 1580 m, Andre 
3640 (K, NY); near Chiriboga, 1900-2000 m, Fagerlind & Wibom 1932 (S). Santiago-Zamora: 
between Rio Sabanilla and Cannilones Tambo, 1370-2135 m, Camp E 63 (BM); near Mendez, Camp 
E4941 (BM); between Rio Sordo and la Esperanza, 2000-2600 m, Acosta Solis 7353 (BM). 
Tungurahua: Asplund 1 7400 (S); Hacienda Rio Verde Grande, 1 500 m, Asplund 7857 (S). 

Geographical range: confined to Ecuador. 

129. Selaginella sulcata (Desv.) Spring ex Martius in Flora, Jena (Beibl.) 1837 (2): 126 
(1837); Baker, Fern Allies : 63 (1887), excl. syn. A. Braun; Knox in Trans, bot. Soc. Edinb. 
35 : 286 (1950). Lectotype from Brazil: without exact locality, in herb Desvaux (P); apud. 
P. Somers, pers comm. 

Lycopodium sulcatum Desv. ex Poiret End. Suppl. 3 : 549 (1814); Sprengel, Syst. Veg. 4:19 

(1827). Lectotype as above. 
Lycopodium stoloniferum sensu Raddi, PL Bras. Nov. Gen.: 81 (1825), non Sw. (1788). 
Selaginella lorentzii Hieron. in Engl., Bot. Jahrb. 22 : 419 (1896). Type from Argentina. 
Selaginella sulcata var. cruenta Martius in Flora, Jena (Beibl.) 1837 (2) : 126 (1837). Type 

from Brazil. 
Selaginella rubescens Hieron. in Hedwigia 43 : 236 (1904). Syntypes from Brazil. 
Selaginella caudorrhiza Baker in J. Bot, Lond. 21 : 334 (1883), Fern Allies: 59 (1887); 

Alston in Pulle, Fl. Surin. 1 : 167 (1938); Knox in Trans, bot. Soc. Edinb. 35 : 284 (1950). 

Type from Surinam. 
Selaginella biauriculata Hieron. in Engl. & Prantl, Nat. Pflanzenf. 1 (4) : 71 1 (1901). Type 

from Brazil. 
Selaginella sulcata var. biauriculata Hieron. ex Bonap., Notes Pterid. 2 : 140 (1915). Type 

from Brazil. 

Specimens seen: 

SURINAM. Without exact locality: Hostmann 3 (BM, K, type of S. caudorrhiza Baker). 
BOLIVIA. Cochabamba: Antahuacana, Espiritu Santo, N of Cochabamba, 750 m, Buchtien s.n. 
(Rosenstock exsicc. 97: BM), Buchtien, 2253 (US); Espiritu Santo, near Cochabamba, Bang 1287 
(BM, NY, US). La Paz: Hacienda Simaco, Tipuani, 1400 m, Buchtien 5281 (US), 5282 (BM, US), 5285 
(BM, NY, US); Bopi River, 915m, Rusby 392 (NY, US); Huachi, Beni River, 1 800 m, White 540 (NY, 
US); Tumupasa, 550 m, Williams 1405 (NY, US); Polo-Polo, near Corvico, Buchtien s.n. (BM), 3556 
(BM); Mapiri, Larecaja, 1525 m, Rusbv 462 (NY). Santa Cruz: Buena Vista, Sara province, 400 m, 
Steinbach 5522 (BM, NY, US), 2068 (BM). 

BRAZIL. Without exact locality : type as above; Swainson s.n. (K); Lobb s.n. (BM, K); Glaziou 46 1 (C), 
2277 (C), 3544 (C), 4490 (BM, C, K, P), 4502 (B), 4503 (C), 5639 (P), 7302 (C), 7356 (C), 7966 (C, P), 
13366 (C, P), 14415 (C, P); Burchell 2238 (K); Sellow 73b (BM); Bowie & Cunningham s.n. (BM); 
Barclay 142 (BM). Amazonas: Glaziou s.n. (BM). Bahia: Blanchet 2207 (P). Ceara: Ubuassu, Dias s.n. 
(BM); Baturite, Santa Clam, Leite 1359 (BM). Espirito Santo: Caparao, Schwacke s.n. (BM); Serra de 
Caparao, Schwacke s.n. (BM); Jatiboca, Brade, Altamiro & Apparicio 18445 (BM); Cachoeiro do 
Itapemirim Faz, Ito. Antonio da Pedra Branca, Brade 19371 (BM). Matto Grosso: Guia, Lindman A 
3493 (NY); H. Smith 147 (BM). MinasGeraes: Fazenda de Grama, Carangoladistr., Mexia 4263 (BM); 
St Hilaire 1430 (P); Angra ds Reis, Hoehne 17417 (BM); Serra do Angico, Fazenda do Diamante, 
Mexia 5644 (BM). Paraiba: Areia, Vasconcellos 260 (BM). Parana: Serra do Mar, Dusen 3797 (BM, P), 
15243 (C), 14375 (C). 609a (C); Rio Branco, Dusen s.n. (BM); Lucena, Wielewski s.n. (NY), 3 (NY), 1 1 



322 A. H. G. ALSTON, A. C. JERMY & J. M. RANKIN 

(BM); San Mattheus, Gansly'\5 (NY); Sao Joao, 545 m, Dusen 1 5423 (BM); Iguazu, Beetle 2054 (BM). 
Pernambuco: Pickel 60 (S); Tapera, Piekel 55 (BM, US). Rio Janeiro: Gaudichaud s.n. (K); Glaziou s.n. 
(BM); Forbes s.n. (BM); Gardner 65 (BM); Mertens s.n. (BM); Du Parquier s.n. (BM); Gaudichaud 135 
(C); Leprieur 199 (C); Mosen 52 (C); Warming 1863 (C); Regnell 298 (C); Corcovado, Martins 197 (M); 
KaflW/ s.n. (B); Jameson s.n. (B, BM); Glaziou 4490 (BM); £ra<te s.n. (BM); Serra Estrella, near 
Mandiocca, Riedel s.n. (K); Nova Friburgo, Glaziou 17961 (C, P, type of 5. biauriculata Hieron.) 
Tijuca, Estrada D. Castorina (Gavea) a vista Chinesa, Alston & Lutz 347 (BM). Districto Federal: 
Lutz 1650 (BM); Sugar Loaf Hill at entrance to harbour of Rio de Janeiro, Jameson s.n. (BM); 
Teresopolis, Sampaio 2688 (BM); Averuda Niemeyer, Freire & Vidal s.n. (BM); Mattas do Rio 
Frapicherro, Peckolt & Freire 617 (BM); Gavea, Frazao s.n. (BM); Nictheroy, Frubuky, Brade 12833 
(BM), 15717 (BM). Rio Grande do Sul: Irai ad flumen Uruguay superius, Emrich s.n. (BM); Taquara, 
Caracol, Dutra 1611 (BM); Porto Novo, ad ripam fluminis Uruguay, Dutra 687 (BM). Santa Catarina: 
Joinville, Schmalz 152 (NY); Mueller 138 (BM, NY); Lages, Spannagel 13 (NY); Itapiranga-Chapeco, 
Reitz 3861 (BM); Hapiranga, ad fluminis Uruguay-Peperi, Rambo 49819 (BM). Sao Paulo: Campinas, 
Novaes 7418 (BM); Ypiranga, Luederwaldt 1659 (BM), 7644 (BM); Guaruja, Santos, Usteh 21024 (S), 
2 1 032 (S); Cantareira, Brade 5138 (NY); Iguape, Morro das Pedra, Brade 15718 (BM). 

Geographical range: Surinam and French Guiana (Somers, pers. comm.), Bolivia to south 
Brazil, Argentina and Paraguay. 

Notes: S. sulcata is distinguished from all the other prostrate species with articulate stems by 
the two long straight eciliate auricles of the axillary leaves. The lateral leaves have long 
finger-like, eciliate auricles on the upper side. The auricles in S. diffusa (No. 133) are more 
deflexed and usually long-ciliate. 

130. Selaginella horizontalis (C. Presl) Spring in Bull. Acad. r. Belg. 10:226 (1843), in 
Mem. Acad. r. Sci. Lett. Belg. 24 : 264 (1850); A. Braun in Annls Sci. nat. (Bot.) V, 3 : 292 
(1865), in Mber. K. preuss, Akad. Wiss. 1865 : 187, 194 (1866); Knox in Trans, bot Soc. 
Edinb. 35 : 285 (1950); Alston in J. Bot., Lond. 72 : 226 (1934). Type from Peru, Haenke 
s.n. (PR). 

Lycopodium horizontale C. Presl, Rel. Haenke : 78 (1825). Type as above. 

Selaginella suavis var. haenkeana Spring in Flora, Jena 21 : 197 (1838). Type from Peru. 

Selaginella fendleri Baker in J. Bot, Lond. 21 : 334 (1883), Fern Allies : 59 (1887). Type 
from Panama. 

Specimens seen: 

VENEZUELA. Apure: Llanos de l'Apure, Geay s.n. (P). Aragua: San Juan de los Morros, 400 m, 
Alston 6034 (BM), 6050 (BM). Carabobo: Carobobo, 500 m, Alston 5606 (BM); Cojedes: near San 
Carlos, Alston 6306 (BM); Aguarita, between Tinaquillo and San Carlos, Alston 6319 (BM); near 
Acarigua, Alston 6320 (BM). Zulia: near El Vigia, Rio Chama, Vareschi & Pannier 1 653 (BM). 
COLOMBIA. Antioquia: Mulatos, 50 m, Haught 4869 (BM, C); Rio Ampurrumiado, 200 m, Gutierrez 
& Barkley 17c 199 (S). Cauca: Carretera a la Plata, de Purace al Alto de San Rafael, 2660-3450 m, 
Garcia- Barriga & Hawkes 12869 (COL). Choco: Saltos de Truando, Schott s.n. (BM, NY, US); 
Quibdo, Rio Atrato, 60 m, Archer 1805 (BM); Ciudad Mutis (Bahia Solano), 5 m, Pinto & Kotschwar 
131 (COL); Negria, c. 50 m, Killip 35023 (COL); between Quebrada Guarapo and Mandinga, 
120-180 m, Killip 35197 (COL). Magdalena: Cano Blanco, SE of La Jagua, foot of Sierra Perija, 
200 m, Haught 3629 (BM); Santa Marta, von Rohr 89 (BM); Agua Dulce Road, Santa Marta, 
305-1065 m. Smith 2238 (BM, NY, US); Don Diego, near sea level, Smith 2699 (NY, US); Rio Frio, 
Santa Marta, O'Neill 1245 (US). Meta: Guatiquia, near Villavicencio, 500 m, Alston 7592 (BM); near 
El Mico Airstrip Rio Guejar, 400 m, Philipson, Idrobo & Fernandez 1373 (BM). Putumayo: Mocoa, 
550-730 m. Schultes & Cabrera 19062 (US); near Puerto Asis, 375 m, Ewan 16759 (BM). Valle del 
Cauca: bridge over Dagua, near Buenaventura, Alston 8610 (BM); Buenaventura, Wall s.n. (S); Rio 
Dagua, near Cordoba, Killip 5045 (NY, US), 5049 (US); Santa Rosa to Villa Cisneros, 250-350 m, 
Killip 5365 (US). 

ECUADOR. Pastaza: Mera, along Rio San Jorge and Rio Tigre, 1200 m, Ollgaard & Balslev 9203 
(AAU,BM). 

Geographical range: Costa Rica southwards to Peru. 



SELAGINELLA IN TROPICAL SOUTH AMERICA 323 

Notes: S. horizontalis is most readily separated from S. humboldtiana (No. 132) by the long 
cilia on the auricles of the lateral leaves. In the dry season contracted stems with crowded 
thicker leaves are produced at the tips of normal stems. The apices of the stem are often 
ascending which is apparent from dried material by the descending rhizophores. 

131. Selaginella microtus A. Braun in Annls Sci. nat. (Bot.) V, 3 : 293 (1865). Type from 
Ecuador, Guayas: Balao, Guyaquil, Jameson 374 (BM, G). 

Selaginella lingulata sensu Baker, Fern Allies : (1887), p.p. non Spring (1 850). 

Specimens seen: 

COLOMBIA. Meta: Puerto Lopez, intendencia del Meta, 240 m, Little 8389 (BM). Valle del Cauca: 

bridge over Rio Dagua, near Buenaventura, Alston 8658 (BM). 

ECUADOR. Without exact locality: Fraser s.n. (BM). Guayas: type as above; Guyaquil, Hall s.n. (K); 

between the airport and textile factory San Antonio, Guyaquil, Asplund 2041 3 (S). Los Rios: Hacienda 

Clementina, Rio Pita, Asplund 5264 (S), 5332 (S); Junction of provinces Guayas, Canar, Chimborazo 

and Bolivar: foothills of W Cordillera, near Bucay, 305^60 m, Camp E3709 (BM). 

Geographical range: Colombia and Ecuador. 

Notes: The median leaves of the main stem in S. microtus are broadest above the middle, 
abruptly acuminate at the apex, and constricted at the base of the shortly ciliate auricle, 
which is half as long as the lamina and peltate. In S. diffusa (No. 133) the median leaves are 
broadest about the middle, gradually acuminate, at the apex, and less constricted at the base 
of the long-ciliate, but smaller, auricle. The median leaves of S. microtus normally show two 
unequal auricles. Moreover, the lateral leaves appear as if attached to the lower surface of the 
main stem, whereas in S. microtus they are inserted laterally. 

132. Selaginella humboldtiana A. Braun in Annls Sci. nat. (Bot.) V, 3 : 293 (1865). Type 
from Venezuela, without exact locality: Orinoco, Humboldt s.n. (B, fragment at BM). 

Selaginella sulcata var. cruenta sensu Baker, Fern Allies : 63 (1887), p.p. quoad syn. A. 
Braun. 

Specimens seen: 

VENEZUELA. Without exact locality: type as above. 

COLOMBIA. Antioquia: Virginias, 40 km E of Puerto Berrio, Archer 1449 (BM, US); Isletas, Rio 

Nare, Andre 436 (NY). Choco: Andagoya, 70-100 m, Killip 35489 (COL). Magdalena: Africa, Sierra 

Perija, 1500 m, Haught 4491 (BM). Norte de Santander: near La Esmeralda, 750 m, Killip & Smith 

20934 (NY). Santander: near Barranca Bermeja, Haught 2014 (BM). Tolima: Mariquita, bosque del 

Acueducto, 550 m, Murill